A simplified key to common genera of terrestrial earthworms

I. Clitellum commencing in front of the 15th segment (not found in the British Isles) 2 - Clitellum commencing after the 15th segment 4 (LUMBRI CIDAE)

2. Setal arrangement perichaetine (Fig. 2) Pheretima (MEGASCo• LECIDAE) - Setal arrangement lumbricine 3 3. Male pores on segment 17 or in 17/18 Spermathecae posterior to segment 10 Eudrilius (EUDRI• LIDAE) Male pores on segment 18 Spermathecae anterior to segment 10 Diplocardia (ACANTHo• DRILIDAE) Male pores on segment 19 (semi-aquatic) Spargano• philus (SPARGANo• PHILIDAE) 4. Prostomium tanylobous (Fig. I), setae closely paired (Fig. 2) at least over part of the body Lumbricus - Prostomium epilobous (Fig. I), or iftanylobous, setae widely paired or distant over the whole body 5 246 BIOLOGY OF EARTHWORMS 5. Clitellum ends after segment 28 6 - Clitellum ends before segment 28, body section quadrangular Eiseniella 6. Tubercula pubertatis absent, or exceptionally, present as simple thickenings of the edges of the c1itellum Bimastos - Tubercula pubertatis present as ridges or isolated papillae (Plate 5b) 7 7. Setae widely-paired or distant, at least posteriorly 8 - Setae closely-paired throughout the length of the body 9 8. Setae widely paired or distant throughout the length of the body, tubercula pubertatis as ridges (excep• tionally as separate tubercles) extending over only part of the length of the c1itellum Dendrobaena Setae closely-paired anteriorly (in region of the hearts), distant posteriorly. Tubercula pubertatis as ridges as long or'longer than the c1itellum Octo/asion 9. Spermathecal pores (Fig. 3) in line WIth seta 'd' or more often near the mid-dorsal line. Body trapezoidal in cross-section. Eisenia and Eisenoides Spermathecal pores situated laterally between setae 'c' and 'd' or 'a' and 'b' and 'c' and 'd'. Body not trapezoidal in cross-section 10 10. Prostomium with longitudinal ridges Eophila - Prostomium without longitudinal ridges II I I. Calciferous glands with two lateral pouches in seg• ment. Terrestrial Allolobo• phora Calciferous glands without lateral pouches. Am- phibious Helodrilus 'Note. Both Sparganophilus and Helodrilus species live in mud bordering streams, rivers and lakes, which should provide confirmation of identi• fication if the key does not provide sufficient data.'

Simple key to species of terrestrial earthworms 'The first part of the following key describes species belonging to families other than the Lumbricidae. The great majority of the repre• sentatives of these families are to be found as indigenous and peregrine species throughout tropical and subtropical parts of Africa, South America and Asia. However, certain members of these families are to SIMPLIFIED KEY TO COMMON GENERA 247 be found in Europe, and particularly )J"orth America, usually as pere• grine species (but occasionally indigenous, such as Diplocardia spp.). The commoner species found in North America are listed.'

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

ACANTHODRILIDAE Genus Dlplocardw I. Clitellum forms a complete ring around the body. 40--120 DiplocardIa mm. 90-120 segs. Anterior dorsal surface pale flesh coloured. singulctris (Ude) Clitcllum 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. I25-160 segs. Anterior dorsal surface (Garman) pale flesh coloured. Two pairs of spermathecal pores in segmental grooves 7/8 and D. riparza 8/9. 200-270 mm. 135-160 segs. Aflterior dorsal surface dark (Smith) brown. 248 BIOLOGY OF EARTHWORMS

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

SP ARGANOPHILIDAE Dorsal pores lacking. Prostomium zygolobous (Fig. I). Anus Spargano- dorsal. phi/us eiseni (Smith) The species of the Family Lumbricidae tabulated here are those which are found most commonly in Britain, North America and in Europe. Amongst these species will be found those that have been trans• ported to various other parts of the world. British and American species are indicated. Although misidentification may arise on the basis of the characters in the first three columns if species not in the key are found, reference to the additional descriptions given in the fourth column should usually be sufficient to avoid confusion.

LUMBRICIDAE Genus Lumbricus Tubercula [st dorsal Clitellum pubertatis pore 26,27-32 28-3 1 7/8 Red/brown or red/violet, Lumbricus irridescent dorsally, pale rubellus·t yellow ventrally. 25-105 Hoffmeister mm. 95-120 segs. 28-33 6/7 Chestnut to violet brown; L. cas• brown/yellow ventrally, taneus. t irridescent, c1itellum (Savigny) orange. 30-70 mm. 82- 100 segs. 31, 32-37 7/8 Setae widely-paired both L. terres• ends of the body, strongly (ris. t pigmented, brown-red Linnaeus dorsally, yellowish ven• trally. 90-300 mm. 110- 160 segs. .t 5/6 Red-brown, lighter ven• L. festivus trally, irridescent dor• (Savigny) sally, prominent c1itel• lum. Not found in large numbers. 48-108 mm. 100-143 segs.

t North American species. .. British species. SIMPLIFIED KEY TO COMMON GENERA 249

Tubercula I st dorsal Clttellum pubertatis pore Genus Eiseniella 22, 23-26, 23-2S, 26 4/S Male pores in 13. Dark Eiseniella 27 brown, greenish, golden tetraeda f yellow, red. Body quad• typica* rangular behind the c1i• (Savigny) tellum. 30-60 mm. 60- 90 segs. Male pore in IS. Other• E. tetraeda " " wise as f. typica (Savigny) f hercynia* (Michaelsen) Genus Bimastos 20, 21, Absent Setae ab > cd. Red• Bimastos 22-29,30 brown. gieselerit 105-1 I S segs. s0--80 mm. (Smith) 22-29 S/6 Reddish-brown, B. tumidi/ segs 30-60. Eisen 20-S0 mm. 23-28 Absent S/6, 6/7 20-S0 mm. 40--60 segs. B. palustrist up to 7S mm. (Moore) 23,24, 24,2S S/6 Reddish dorsally, yel- B. parvust 2S-3 I , 32 26-30 lowish ventrally. 2S-40 (Eisen) or absent mm. 90--110 segs. Absent 5/6 Rose-red. B. longi• 98-122 segs. 60--<)0 mm. cinctus t (Smith & Gittins)

24, 2S-32, Absent S/6 Prostomium tanylobous, B. eiseni* 33 body cylindrical, reddish (Levinsen) or violet dorsally, yel• lowish ventrally, c1itel- lum red. 30-64 mm. 7S- I I I segs. Absent Pale red to chestnut B. zeteki t brown, often localized (Smith & whitish banding anter- Gittins) iorly. 100--140 mm. 110- 140 segs. None S/6 Dark reddish brown to B. heim- brown setae closely burgen ·t paired. (Smith) 250 BIOLOGY OF EARTHWORMS

Tubercula I st dorsal Clitellum pubertatis pore

Genus Dendrobaena 25, 26-31, 28-30 5/6 Rosy to deep red, last Dendrobaena 32 posterior segments yel~ rubida low, 27-90 mm. 50-100 (Savigny)* segs. .r subrubi• cunda (Eisen) 25, 26, 27- 28, 29-30, Setae widely paired, dor- D. rubida 30, 31, 31 or sally red-brown with (Savigny)* 32, 33 absent bluish tint. Intersegmen- .r tenuis tal grooves and ventral (Eisen) side light. 20-85 mm. 90-120 segs. 25,26-28, 5/6 or Smoky-grey with red D. platyura 29,30 sometimes pigment posteriorly. 80- (Fitzinger) 8/9 180 mm. 120-160 segs. 26, 27-31, 5/6 Dark red dorsally, lighter D. rubida*t 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 .r typica* bands separated by non- (Rosa) pigmented zones. Some- times uniformly unpig- mented. 50-155 mm. 80-- 225 segs.

31-32, 33 4/5 Red, violet, yellow or D. copper. Posterior octa- octaedra*t gonal. 17-40 mm. 79-95 (Savigny) segs. 28-33, 34 5/6 Dorsally pale red, 1st D. attemsl segment and ventrally (Michael• and clitellum white. 20- sen) 50 mm. 100--150 segs.

4/5 Red-violet, slightly irri- D. descent. 30-65 mm. 83- mammalis*t 100 segs. (Savigny) Undetect- Red dorsally or unpig- D. pygmaea* able mented. 30-32 mm. 103- (Savigny) 180 segs. SIMPLIFIED KEY TO COMMON GENERA 251

Tubercula 1St dorsal Clztellum pubertatls pore Genus Octolasium 11/12 Blue-grey with (usually) Octolasion lilac-blue dorsal line. cyaneum*t Last 4~5 segments yel• (Savigny) low, anterior segments pink, clitellum red• orange. 50~I60 mm. IOO~

~I50 segs. 8/9, 9/10 White, grey, blue or O. lacteum*t or 101 II rose-pink, clitellum pink (Oerley) or orange. 30~I60 mm. 90~I80 segs.

Genus Eisenia 24, 25, 28~30, 31 4/5 Red, purple or brown; Eisenia 26--J2 yellowish ventrally. Dor• !oetida*t sal surface pigment alter• (Savigny) nating with light inter• segmental zones. 32~I30 mm. 80~I 10 segs. 27, 28~32, 30-32 5/6 Light to bright reddish, E. 33,34 mostly on dorsal aspect. hortensis* t (Michael• sen) Genus Eisenoides 24, 25~30, 27~39 5/6 Reddish, body cylindri- Eisenoides 31 cal. 95~I20 mm. I03~I58 carolin• segs. Setae closely paired. ensls t (Michael• sen)

4/5 Pigmented brown never E. usually blood shows lonnbergit 90~I75 mm. 100~152 (Michael- segs. sen) Genus Eophila 33, 34 35, 36, Yellowish or grey. 52~ Eophila 140 mm. 132~I70 segs. lcterica* (Savigny) Genus Allolobophora 25, 26~31, 29~30, 31 4/5 Pale red, without pig- Allolo• 32, 33 or 30~32 ment, prominent and bophora flattened clitellum. 25~ rosea*t 85 mm. 120~150 segs. (Savigny) 252 BIOLOGY OF EARTHWORMS Tubercula ISt dorsal Clitellum pubertatis pore

26-32 28, t29 5/6 Grey, unpigmented, A. culpifera and t29, 30 usually blood shows (Tetry) through epidermis. 20- 60 mm. 80-120 segs.

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

26,27, 31 and 33 rr/12 Anterior, especially the A. caliginosa 28-34,35 or 12/13 first few segments pink, f typica*t otherwise pale grey, yel- (Savigny) lowish clitellum. 40-100 mm. 120-150 segs.

26,27 31-33 Colour as for f. typica. A. caliginosa 28-34, 35 Tubercula pubertatis of (Savigny) two raised tubercles, f tra- connected by a narrow pezOl des t bridge. (Duges)

27-32,33 Absent Indistinct Grey, unpigmented. 80- A. miniscula 100 segs. 22-25 mm. (Rosa) 27, 28-35 32 and 34 Body cylindrical, pale A. terrestris grey, unpigmented. 90- (Savigny) 150 mm. 160-200 segs.

27, 28-35 32-34 Body cylindrical, colour A. longa*t as for A. terrestris. Ude 90- 150 mm. 17 1- 181 segs. 27, 28-35 31 and 33 Segments posterior to A. nocturna* (extending 13 divided by two Evans into grooves into three rings segment 32) Dark reddish brown, cli- tellum paler. 90-180 mm. 200-250 segs.

27,28-34, 31-35 10/II 55-100 mm. 152- 194 A. tubercu- 35 II/I2 segs. Body cylindrical, culata* t or 12/13 unpigmented, greyish. Eisen 28-35, 36 33 and 34 4/5 or 5/6 Unpigmented, anterior A.limicola*t pink, the rest of the body Michaelsen pinkish grey. Bulbous anterior. 40- 100 mm. 86-146 segs. SIMPLIFIED KEY TO COMMON GENERA 253

Tubercula IS! dorsal Clitellum Puber!atis hore

31 and 33 4/5 Light or dark green, yel- A. chlor• and 35 low, grey, pink, slate- otica·t blue, c1itellum pink, (Savigny) 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- ISO segs.

Nomenclature Allolobophora caliginosa Cernosvitov and Evans (1947) described Allolobophora caliginosa as two varieties, f. typica Savigny and f. trapezoides Duges, depending upon whether the papillae of the tubercula pubertatis were distinct, or joined by a narrow translucent bridge. Cain (1955) considered that all British specimens should be called A. caliginosa Savigny, and the name A. trapezoides Duges be given to continental specimens which, apart from the narrow bridge, have other characters which separate them from A. caliginosa, such as differences in the number of segments, position of the first dorsal pore and genital markings, and extension of the glan• dular tumescences around the male pores. Although Omodeo (1956) called the continental worms A. caliginosa f. trapezoides, Gerard in his synopses of the British Lumbricidae (1964) agreed with Cain, and listed only A. caliginosa Savigny, with no varieties. Gates (1972) also recog• nized A. trapezoides as a separate species. At the same time, Gates 254 BIOLOGY OF EARTHWORMS postulated that the specific name cafiginosa for the British species is in• correct, and on the evidence of original identification by Eisen of a group of Scandinavian earthworms, and identifications by Friend (1923) and Davies (1960) states that the correct name should be turgida. Gerard disagrees on the grounds of priority. Nevertheless, Allolobophora turgida Eisen is used frequently, especially by American authors in place of Allolobophora cafiginosa Savigny. No doubt, in the next few years, this dichotomy will be resolved.

Octolasion lacteum Oerley Very close indeed in relationship to O. cyaneum, almost indistinguishable except for a one segment difference in position of clitellum and tubercula pubertatis, it has been suggested (Omodeo, 1959) that they are diverse forms of the same agamous species, i.e. parthenogenetic morphs. The name given to this 'species', lacteum in 1881 is in common use, but Savigny's Enterion tyrtaeum 1826 was recognized by Cernosvitov as the same species (as lacteum) although he merely left it as a foot-note instead of claiming priority for tyrtaeum over lacteum. Gates (1972) has reinvesti• gated the situation, and states that the name of this taxon must be Octolasion tyrtaeum Eisen. Both tyrtaeum and lacteum are currently used in literature. Because of the many changes in taxonomic nomenclature that have taken place during recent years, many authors continue to use names which have since been superseded, or which are still well-established. A list of such synonyms is given below. Eisenia rosea Savigny = Allolobophora rose a Savigny. Allolobophora minima Muldal = Allolobophora muldali Omodeo. Bimastos beddardi Michaelsen = Bimastos parvus Eisen.

Eisenia veneta. Rosa = Dendrobaena veneta f. typica Rosa. Blmastos tenuls Eisen = Dendrobaena rubida Savigny f. tenuis Eisen. Allolobophora icterica Savigny = Eophlla ieterica Hoffmeister. Eophila oculata Hoffmeister = Helodrilus oculatus Hoffmeister. Dendrobaena subrubicunda Eisen = Dendrobaena rubida Savigny f. Sub- rubicunda Eisen. Genl,ls Octo/asion oerley = Octo/asium oerley. SIMPLIFIED KEY TO COMMON GENERA 255 The following species, otherwise placed in the genus Allolobophora Eisen, will also be found (especially in current North American publi• cations) in the proposed new genus Aporrectodia Gates: lzmlcola, longa, moebii, rosell, trapezoides, tllberculata, tllrgida. Glossary

Acanthodriline. Having the male pores in segment 18 and the prostatic pores in segments I7 and 19. Aclitellate. Without a clitellum. Not necessarily confined to juvenile stages. Amphimitic. Sexual reproduction in earthworms involving two parents. Anthropochorous. Transported by the agency of man (usually unin• tentional). Astomate. The condition in which the nephridia are closed, i.e. without a nephrostome. Autotomy. The process of self induced loss of segments - as when a worm is gripped by the tail. Brown bodies. Rounded bodies containing tissue debris such as setae and corpuscles, and foreign material. Chaeta, chaetae. Alternative for seta, setae. Classical system. The classification of the Oligochaeta originated by Michaelsen in 1900, and developed by Stephenson in 1930 in 'The o ligochaeta' . Clitellate. Possessing a clitellum. Also used to describe the stage in a worm's growth during which it has a clitellum, when it is a more specific term than 'adult'. Clitellum. A region of epidermal thickening, containing gland cells which secrete the cocoon material. Copulatory chamber. An invagination of the body wall into the coelom which contains the male pore. Copulatory pouches. An old term for spermathecae. Copulatory setae. Setae in the same segment as, and near to, the spermathecae. Diapause. As applied to earthworms, a non-active state during which the worm has an empty gut and stays tightly coiled in a mucus lined 258 BIOLOGY OF EARTHWORMS cell within the soil to protect itself against adverse environmental conditions. It may be optional or obligatory. Endemic. Indigenous. Epilobous. The condition when the continuation of the prostomium q. v. backwards does not reach the first segmental groove. Estivation. A summer diapause. Genital tumescences. Raised areas of the epidermis from which genital setae grow, found in the Lumbricidae. Haemerophilic. Not adversely affected by human interference with the environment. Haemerophobic. Adversely affected by human interference with the environment. Heteromorphic. The condition when regeneration occurs so that a head regenerates instead of a tail, and vice versa. Holandric. Testes restricted to segments 10 and II, or a homeotic equivalent. Hologynous. Ovaries restricted to segments 12 and 13, or a homeotic equivalent. Homeotic. The state in which an organ or series of organs are in a seg• ment or segments in which they are not normally found. Intersegmental furrow. The actual boundary between two adjacent seg• ments. Intersegmental groove. The visible annular depression that indicates externally the position of the boundary between two adjacent seg• ments. Juvenile. The term used for earthworms from the time of emergence from the cocoon to when the first indications of maturity, such as genital tumescences, seminal grooves and genital pores appear. Lumbricine setal arrangement. Having four pairs of setae per segment. Macroic. Large. A substitute for meganephridial. Male ducts. Male gonoducts. Sperm ducts q.v. Male funnels. Funnel shaped internal end of sperm duct. Male pores. Exterior openings of the sperm ducts. Meganephridia. Large nephridia. A term now little used, the condition meganephridial now being encompassed by the term holonephric. Micronephridia. Small nephridia, usually numerous, often encompassed by the term meronephric. Meronephridia. Divided nephridia, individual tubules often present in large numbers. Can be either large - megameronephridia, or small - micro meronephridia. GLOSSARY 259

Megascolecoid. Referring to worms placed III the classical family Megascolecidae. Oviducts. Female gonoducts. Parthenogenesis. Reproduction in which the ova develops without being fertilized by a spermatozoa, hence it only involves one parent. Peregrine. Foreign, not indigenous. The agency is almost always trans• portation by man. Perichaetine setal arrangement. Many setae arranged in a ring right round each segment with only a break in the mid-dorsal and mid• ventral regions. Peristomium. The most anterior segment of an earthworm surrounding the mouth, and which bears the prostomium. Phylogeny, Phylogenetic. Pertaining to past evolutionary development as opposed to individual development (autogeny). Polyandric. Having testes in more than segments 10-1 I. Polymorphic. Pertaining to polymorphism. Polymorphs anslllg from parthenogenesis and geographical distribution are considered important in systematics. Proandry, Proandric. Testes restricted to segment 10 or a homeotic equivalent. Progyny, Progynous. Ovaries restricted to segment 12, or a homoeotic equivalent. Prolobous. The condition when there is no continuation of the pros• tomium backwards into the peristomium. Prostate. Glands associated with the male ducts, usually tubular, open• ing beside the male pores in acanthodrilid, octochaetid and ocnero• drilid worms. Prostomium. A lobe-like appendage attached to the front of the dorsal aspect of the peristomium. Racemose. Applied to prostates when those organs are divided into many lobes (cf. a bunch of grapes). Seminal grooves. Permanent longitudinal grooves on the ventral surface of an earthworm associated with the male and prostatic pores which form a passage for seminal and other fluids. Seminal vesicles. Septal pockets in which spermatozoa mature. Seta. Setae. Stiff bristle-like structures protruding from follicles through the epidermis. Usually sigmoid in shape, except specialized setae. May be enlarged at the extreme portions of the body. Somatic. Pertaining to any part of the body except the genital organs. Sperm ducts. Ducts carrying sperm to the bodies exterior. 260 BIOLOGY OF EARTHWORMS Stomate. Pertaining to an 'open' nephridium, i.e. a nephridium with a funnel opening (usually) to the exterior. Tanylobous. The condition when the continuation of the prostomium reaches backwards to the first segmental groove. Taxon. Any unit in a system of classification. Testis sac. Closed part of the coelom containing the testes and male funnels of a particular segment. Zygolobous. Condition when the prostomium as seen from above, is not differentiated from the peristomium by any superficial markings. References

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

Acanthodrilacae, 44 dubiosa, 226 Acanthodrilidae, 44, 46,51-2, 61, 65, 66, japonica, 137 67, 74, 75, 122, 245, 247 limlcola, 62, 241, 252 Acanthodrilinae, 43, 44, 45, 46 longa, 62, 69, 70, 71, 73, 80, 84, 93, 108, Acanthodrilini, 45 II I, 128, 130, 131, 138, 140, 145, 146, Acanthodrilus, 41, 51 150, 153, 154, 161, 162, 165, 172, 173, Actinomyces, 187 192, 193, 199,216,224,227,231,232, advenltlum, Blpalium, 147 234, 240, 2... , 252 Aeolosomatidae, 13, 4 I, 42 minzma, 256 aerophilus, Thomznx, 227 mlnzscula, 252 affinis, Dichogaster, 80 muldall, 62, 256 africanus, Hyperiodrilus, 75, 136, 142, 143, noclurna, 69, 70,84,87,125,126, 12M, 129, ISO, 154, 157 130, 131, 132, 134, 136, 13M, 140, 145, afrlcanus, Ocnerodrilus, 226 149, ISO, 151, 161, 162,232,234,240, Agastrodilus, 52 241, 252 agglutinatus, Bacdlus, 185 robusl a, 146 agItans, Uropoda, 149 rosea, 9, 63, 70, 71, 73, 81, 83, 125, 126, agrestis, Pheretlma, 63, 107 127, 128, 129, 130, 131, 132, 133, 134, Aikinetocystzs, 148 138, 145, 146, 150, 151, 152, 157, 161, alexandrt, Pheretlma, 79 165, 216, 231, 232, 239, 240, 251, 256 Allolobophora, 59, 69, 81, 82, 127,228,246, terrestns, 146, 252 250, 251, 25 6 terrestris f. longa, 73 antipae, 30, 252 trapezoldes, 62, 255 caligznosa, 26, 68, 70, 71, 73, 80, 83, 87, tuberculata, 62, 80, 93, 241, 252 98, 102, 117, 125, 126, 128, 129, 130, turglda, 62, 80, 255 13 1, 132, 133, 134, 135, 136, 138, 139, Alluroididae, 43 145, 146, 149, ISO, IS I, 152, 153, 154, Alma, 57, 75, 157, 158 157, 158, 161, 162, 163, 164, 165, 167, Amoebotaenza cuneatus, 226 172, 174, 176, 179, I!!9, 197, 19/5, 200, lumbri, i, 226 201, 216, 226, 227, 229, 232, 239, 240, Andiorrhinus, 56 241, 255 AndlOswlex, 56 caligznosa f. tYPlca, 63, 252, 255 annulata, Capil/arza, 226 caligznosa f. trapezoides, 26, 63, 80, 252, anomala, Mil/soma, 74, 125, 136, 138, 152 255 anomala, Pheretima, 79 chlorolica, 62, 68, 70, 71, 72, 73, 80, 83, Anoplophyra, 148 93, I I I, II8, 125, 126, 128, 129, 130, antipae, Al/olobophora, 30, 252 13 1,133,134,135,136,145,146,148, apricaria, Pluvla/zs, 147 ISO, 151, 152, 153, 154, ISS, 161, 164, Aporrectodza, 256 170,200,216,219,229,231,232,239, /zmlcola, 256 240, 241, 242, 253 longa, 256 culpifera, 252 moebll, 256 302 BIOLOGY OF EARTHWORMS Aporrectodia--cont. Caleb/ella, 52 rosea, 256 (!Iiz/ormco, Pherer;ma, 22, 25, 1,3, 102, 154, trapezoides, 256 157, 158, 247 tuberculata, 256 caligmosa, Allolobophora, 26, 68, 70, 71, 73, turgida, 256 80,83,87,98, 102, "7, 125, 126, 128, Aptodrilus, 56 129, 130, 131, 132, 133, 134, 135, 136, Ascaridia gallt, 228 138,139,145,146,149, ISO, 151, 152, Ascaris suum, 228 153, 154, 157, 158, 161, 162, 163, 164, Aspidodrtlus, 146 165, 167, 172, 174, 176, 179, 189, 197, altemst, Dendrobaena, 170, 239, 250 198, 200, 201, 216, 226, 227, 229, 232, austrabs, Megascolides, 19, 81 239, 240, 241, 255 avestcula, Udema, 79 caltginosa f. trapezOldes, Allolobophora, 26, Azotobacter, 175, 183 63, 80, 252, 255 caltgmosa f. ryptca, Allolobophora, 63, 252, Bactllus agglutinatus, 185 255 botulinus, 148 Calltdnlus, 57 cereus, 182, 185 campanulata, Pherettma, 79 cereus var. mycoides, 184 canus, Larus, 147 tdosus, 185 Captllaria annulata, 226 Balanteodrtlus, 5I causmflata, 227 Barogaster, 52 mucronata, 227 120 Bassia longifolia, plica, 227 beatrix, Octochaetana, 80 putom, 227 beddardi, Btmastos, 62, 150, 256 carnana, Sarcophaga, 149 Beddardiella, 54 carolinensis, Eisenoides, 251 Benhamia, 52 castaneus, Lumbricus, 38, 62, 69, 70, 71, 73, Benhaminae, 44 80,82,93,98, ,,8, 126, 127, 128, 145, bermudensis, Pontodrilus, 61, 79 146, 150, 161, 163, 164, 165, 172, 232, Bettonta, 54 241, 242, 248 btcmta, Pheretima, 63 causinflata, Captllana, 227 Bimastos, 39, 59, 64, 246, 249 Causus rhombeatus, 147 beddardt, 62, ISO, 256 Celenella, 52 elsem, 13, 127, 128, 145, 149, ISO, 159, 163, 165, 170, 240, 241, 245, 249 cereus var. mycoldes, BaCIllus, 184 gieseleri, 62, 249 Cestoda, 225, 226 heimburgeri, 62, 249 Chi/ota, 51 tcenorum, 241 chlorot/ca, Allolobophora, 62, 68, 70, 71, 72, longicinctus, 62, 249 7~80,83,93, III, 118, 125, 12~ 12~ lonnbergi, 149, 150 129, 130, 13 1, 133, 134, 135, 136, 145, muldali, 241 146, 148, 150, 151, 15 2, 153, 154, ISS, palustris, 62, 249 161, 164, 170, 200, 216, 219, 229, parvus, 20, 62, 93, 241, 249, 256 231, 232,239,24°, 241, 242, 253 tenms, 8, 256 ChunlOdnlus, 54 tumidts, 62, 82, 85, 249 Clitellata, 40 weicht, 62 Clostrtdium, 183 zetek!, 62, 72, 74, 75, 85, 86, 88, 89, 145, coeltcolor, Streptomyces, 187 249 Comarodnlus, 49 Bipalium adventttum, 147 commUniS, Diplocardia, 247 kewensts, 143, 147 commumsstma, Pherettma, 93 bishamban, Ramiella, 80 controversa, Tilletta, 188, 224 Borgert/a, 54 coracana, Eleusme, 228 botulmus, Bactllus, 148 corethrurus, Pontoscolex, 61, 66, 79, 228 Branchiobdellidae, 42 Corvus corone, 147 bolam, Dtchogaster, 63, 80 crassum, Porrocaecum, 227 bronch/alis, Cyathostoma, 226, 227 CraterocysttS, 148 Buttnmodrilus, 54 Criodrilidae, 47,58, 61 SYSTEMATIC INDEX 30 3

Criodrilinae, 43 jiortdana, 122 CrlOdrilus, 58, 114 mlsslssipptensis, 89, 122 Criodrilus lacuum, 61, 75, 226 rtparta, 140, 247 culpifera, Allolobophora, 252 slngularis, 61, 247 cuneatus, Amoebotaenia, 226 Diplocardiinae, 43 cuneata, Ta,nia, 148 Dzplotrema, 41, 51 Curgiona, 50 Dzporochaetti, 51 cyaneum, Octolasion, 62, 70, 82, 128, 129, Distichopus, 148 138,145,150,151,161,165,216,232, Dorgiodrilus, 50 239, 240, 241, 25 1, 255 Drawida, 48, 66 Cyathostoma bronchialis, 226, 227 grandis, 140, 146 nagana, 8o papillifer, 80 Decachaetus, 51 rara, 80 Deccania, 50 Drzlocrzus, 57, 66 dejecta, Pheretima, 79 dubzus, Microscolex, 61, 79 Deinodrilus, 67 dubiosa, Allolobophora, 226 Dendrobaena, 14, 59, 81, 84, 246, 250 dunguensis, Schubotziella, 78 attemsi, 170, 239, 250 mammalis, 62, 6cj, 70, 146, 232, 241, 250 octaedra, 62, 80, 98, 127, 128, 13 1, 145, Echinocystis, 148 146, 149, 150, 159, 163, 165, 170,240, egglestoni, Diplocardia, 75, 84 241, 250 Eisenia, 63, 64, 69, 246, 251 platyura, 250 foetida, 13, 17, 19,62,71,73,78, So, 81, pygmaea, 241, 250 82, 85, 92, 93, 95, 99, 100, 108, 115, rub,da, 62, 80, 82,127,131,145,146,149, 116, 147, 148, 149, 153, 155, 156, 157, 150, 165, 170, 232, 239, 240, 250, 256 158, 159, 172, 186, 188, 189, 196,200, rubida f. tenuis, 81, 241, 250, 256 201,226,227, 228, 229, 232, 239, 240, rubida f. subrubicunda, 146, 241, 250, 256 24 1,25 1 subrubicunda, 19, 70, 71, 73, 82, 86, III, hortenslS, 62, 25 I 150, 153, 154, 192,232, 240, 256 lonnbergz, 251 veneta var. hibernica f. dendroidea, 241 rosea, 62, 63, 72, 73, So, 98, 140, 158, veneta var. hibernica f. typica, 146, 241 256 veneta var. hortensis, 241 veneta, 226, 256 veneta f. tYP,ca, 241, 250, 256 zebra, 62 veneta var. zebra, 241 elseni, Bimastos, 13, 127, 128, 145, 149, Desmogaster, 48 150, 159, 163, 165, 170,240,241,245, Diaphrodrilus, 50 249 Dicelis, 148 eiseni, Lumbricus, 62 D,chogaster, 47, 52, 95 eiseni, S parganophilus, 63, 154, 248 bolaui, 63 Eiseniella, 59,81,246, 249 jaculatrtx, 142 tetraedra, 62, 80, 165, 232, 240, 241, 249 sa liens, 63 tetraedra f. hercynza, 241, 249 Dichogastidae, 45 tetraedra f. tYP,ca, 165, 248, 249 Didymogaster, 49 Elsenoides, 59, 64, 246, 250 sylvatlcus, 19 caro/inensis, 25 I dijJringens, Pheretima, 63, So, 122, 247 Eleuszne coracana, 228 D,gaster, 49 elongata, Pheretzma, 79, 228 Dilepus undula, 226 elongatus, Metastrongylus, ;u6 DtnodrtlOldes, 51 Eminoscolex, 54 Dinodrilus, 52 emissartus, Haplotaxis, 62 Dioctophyma, sp., 227 Enantiodrilus, 56 Dionyx, 148 Enchytraeidae, 8, II, 42, 146, 209 DIplocardia, 51, 61, 64, 13 1, 157, 245 ensicaudatum, Porrocaecum, 227 communis, 247 EnterlOn tyrtaeum, 256 egglestont, 75, 84 Eodrilus, 51, 245 304 BIOLOGY OF EARTHWORMS

Eophda, 60, 64, 66, 246, 250 Gatesla, 50 (Allolobophora) icterica, 150, 241, 251, gieseleri, Bimastos, 62, 249 256 glabra, Pheretima, 79 Eophila oculata, 256 Glossoscolecidae, xiv, 4, 23, 42, 43, 47, Ephymodrdus, 55 55-6, 61, 66, 75 encetorum, Turdus, 147 Glossoscolecinae, 8, 36, 43 Erithacus rubecula, 147 Glossoscolex, 56 Eschmchia colt, 184 Glyphidrilus, 57 Estherella, 56 gordiodes, Haplotaxis, 62 Eudlchogaster, 53 GordlOdrilus, 50 Eudmodrdoides, 5 I gracilus, Pontodrilus, 61 Eudrilidae, 42, 43, 46,53-4,61,66,75,78, grandls, Drawida, 140, 146 79, 83, 245, 248 Grayallia, 148 Eudrilinae, 43, 46,54-5 Gregarma, 148 Eudriloides, 54 Eudnlus, 55, 88 eugemae, 61, 66, 142, 248 haemorrhoidalis, Sarcophaga, 149 eugeniae, Eudrilus, 61, 66, 142, 248 Haasema, 55 Eukerria, 50 haltotidea, Testacella, 147 hortensis, 79 Haplodrilus, 50 saltensls, 63 Haplotaxidae, 43, 61, 62 subandina, 79 Haplotaxis, emiSSarlUS, 62 Eupolygaster, 48 forbesi, 62 Eupolytoreutus, 55 gordlOdes, 62 europaea, Talpa, 147 ichthyophagous, 62 europeus, Evinaceus, 147 Hastirogaster, 48 Euscolex, 55 hawayana, Pheretlma, 63, 80, 247 Eutoreutus, 55 heimburgerl, Bimastos, 62, 249 Eutrigaster, 53 Helodrilus, 55, 60, 64, 66, 246 Eutyphoeus, 53, 95 afrlcanus, 150 waltoni, 78, 142 oculatus, 241, 253, 256 Evinaceus europeus, 147 herculeus, Lumbricus, 227 excavatus, Penonyx, 80, 95 Heterakis, 148 eXlgus, Pheretlma, 79 Heterodera rostochlenSls, 224 Enus, 45, 49 Heteroporodnlis, 49 hdgendorfi, Pheretima, 63, 72, 79 Hlppopera, 55 jestil'us, Lumbncus, 62, 82,127,165,232,241, nzgenae, 181 248 Hirudinea, 40, 41 Fimoscolex, 56 Hlstomona murchiel, 148 Fletcherodrtlus, 49 Histomonas, 226 flondana, Diplocardia, 122 H oloscolex, 56 jiJetida, Eisenia, 13, 17, 19, 62, 71, 73, 78, Hoplttophyra, 148 80, 81, 82, 85, 92, 93, 95, 99, 100, 108, Hoplochaetella, 53 115, 116, 147, 148, 149, 153, 155, 156, Hoplochaetina, 53 157, 158, 159, 172, 186, 188, 189, 196, Hormogaster, 58 200, 201, 226, 227, 228, 229, 232, 239, redli f. gigantes, 65 240,241, 251 Hormogastridae, 58, 65 forbesi, Haplotaxls, 62 Hormogastrinae, 43 Fraxinus pennsylvanica, 198 hortensis, Elsenia, 62, 251 Fndericia parasttica, 146 Eukerria, 79 friendl, Lumbricus, 241 houlleti, Pheretlma, 63 Fusarium, 188, 224 Howascolecini, 45 Howascolex, 53 galli, Ascaridia, 228 hupelensis, Pheretlma, 24, 63, 73, 74, 75, Gardullaria, 55 127, 131, 138, 140, 142, 153, 157, 158 SYSTEMATIC INDEX 305

Hypmodrilus, 55, 79 42, 43, 58, 60, 61, 64, 172, 240, 241, afncanus, 75, 136, 142, 143, 150, 154, 245, 246, 248, 255 157 Lumbriculidae, 2, 5, 6, 22, 30, 39, 40, 42, Hyslrlchis triwlor, 226 47, 80, 98, 225 Lumbricus, 4, 14, 17, 19, 22, 25, 26, 30, 33, Icenorum, Bimastos, 241 34, 36, 37, 60, 62, 81, 83, 84, 85, 110, Ichthyophagous, Haplotaxis, 62 139, 245, 248 icterica, Eophila (Allolobophora), ISO, 241, castaneus, 38, 62, 69, 70, 71, 73, So, 82, 251, 256 93, 98, 118, 126, 127, 128, 145, 146, idosus, Bactllus, 185 150, 161, 163, 164, 165, 172,241,242, Ilyogema, 50 248 maequalis, Ventura, 188 eiseni, 62 mfantalis, Pheretima, 79 festivus, 62, 82, 127, 165, 232, 241, 248 Iridodri/us, 55 friendi, 241 hercu/eus, 227 rubellus, 62, 69, 70, 71, 73, 80, 82, 93, 98, jacu/atrix, Dichogaster, 142 108, III, 127, 128, 145, 146, 150, 154, Japomca, Allolobophora, 137 155, 162, 165, 170, 172, 173, 183, 192, 196, 226, 227, 229, 232, 239, 240, 241, KafJania, 55 248 Kerriona, 50 terrestris,6, 13,21,27,39,62,72,76, So, kewensis, Bipalium, 143, 147 81, 82, 83, 84, 85, 86, 88, 89, 90, 91, Kynotus, 57 92, 93, 94, 97, 98, 103, 106, 107, 108, 114, 115, 117, 125, 126, 128, 129, 130, /acteum, Octo/aston, 82, 98, 128, 146, 153, 13 1, 132, 134, 136, 138, 139, 140, 145, 165, 183,228,229,232,241,251,255, 146, 148, 149, 150, 151, 152, 153, 154, 25 6 156, 157, 160, 161, 162, 166, 168, 169, /acuum, Cnodri/us, 61, 75, 226 172, 173, 177, 178, 179, 182, 183, 184, Lampito, 49, 80, 95 185, 188, 192, 195, 196, 198, 199, 200, mani/ii, 80 201,206,216,218, 21 9,226,227,228, Larus canus, 147 229,23°,231,232,234,235,236,237, latus, Protoscolex, 41 239, 240, 241, 242, 248 Legonea,55 Lennogaster, 53 Mahema, 50 Lennoscolex, 49 Malabarta, 50 pumila, 80 Malabarini, 45 Leptodrtlus, 5 I, 54 Malodrtlus, 55 Leucodrilus, 53 mamma lis, Dendrobaena, 62, 69, 70, 146, /evIS, Pherellma, 63 232, 241, 250 Libyodrilus, 54 mamlti, Lampito, 80 /imicola, Allolobophora, 62, 241, 252 Maortdrtlus, 5 I Liodrtits, 50 marcessens, Serratia, 184 /onga, Allolobophora, 62, 69, 71, 73, 80, 84, 93, mariana, Pieea, 198 108, II I, 128, 130, 131, 138, 140, 145, mauget, Testacella, 147 146, 150, 153, 154, 161, 162, 165, 172, Maupasella, 148 173, 192, 193, 199,216,224,227,231, mays, Zea, 228 232, 234, 240, 241, 252 Megachaetma, 54 longa, Aporrectod,a, 256 Megadrili, xv, 42, 43, 61, 65, 66, 79 /ongicinctus, Bimastos, 62, 249 Megascolecidae, 8, 10, 13, 19, 20, 22, 23, longifolta, Bassta, 120 25, 30, 32, 36, 38, 39, 43, 44, 45, 46, /onnbergi, Blmastos, 149, 150 47, 48-50 , 62, 66, 80, 146, 147, 245, liinnbergi, Eisenoides, 62, 251 247 lot'endgel, Pherellma, 63 Megascolecinae, 7, 43, 44, 45 lumbnci, All%bophora, 226 Megasco/ex, 7, 30, 41, 49, 149 /umbncl, Polycercus, 226 Megascoltdes, 13, 53, 61, 199 Lumbricidae, 8, 10, I I, 12, 13, 20, 35, 36, australis, 19, 81 306 BIOLOGY OF EARTHWORMS

Meles meles, 147 Neodrtlus, 52 meles, Meles, 147 Neogaster, 53 merid,ana, Pheretima, 79 Neummaneona, 55 merula, Turdus, 147 ntgerzae, Hippopera, 181 merulae, Syngamus, 227 NocardIa polychromogens, 187 mesomeltnus, Microsaurus, 147 nocturna, Allolobophora, 69, 70, 84, 87, 125, Metadrilus, 54 126, 128, 129, 130, 131, 132, 134, 136, Metaradiophyra, 148 138, 140, 145, 149, 150, 151, 161, 162, Metascolex, 55 232, 234, 240, 241, 252 Metastrongylus, 148, 225 Notoscolex, 49, 142 elongatus, 226 termlticola, 147 pudentotectus, 226 Notykus, 54 salamI, 226 Microchaetidae, 47, 57, 66, 223 oCCldentalts, Ocnerodrilus, 63, 80 Microchaetinae, 43 Ocnerodrilinae, 43, 44, 45, 46 Microchaetus, 57, 142 Ocnerodrilini, 45 Microdrili, 42, 48, 82 Ocnerodrilus, 5I Microsaurus mesomelinus, 147 afrtcanus, 226 MicrosCIJlex, 51 occidentalis, 63, 80 dublUS, 61, 79 octaedra, Dendrobaena, 62, 80, 98, 127, 128, modestus, 228 13 1, 145, 146, 149, 150, 159, 163, 165, phosphoreus, 61 170, 240, 241, 250 millardi, Perionyx, 140 Octochaetidae, 44, 46,52-3, 63, 66, 67, 80 Millsonta, 53 Octochaetinae, 43, 45, 46 anomala, 74, 125, 136, 138, 15 2 Octochaetini, 45 minima, Allolobophora, 256 Octochaetoides, 53 mtnima, Pheretima, 63, 247 Octochaetona beatrtx, 80 mtniscula, Allolobophora, 252 surensts, 80 mississippiensls, DiplocardIa, 89, 122 OClochaelus, 53 modestus, Mlcrosco/ex, 228 Octo/aslon, 60, 69, 166, 173, 219, 246, 251 moebli, Aporrectodia, 256 cyaneum, 62, 70, 82, 128, 129, 138, 145, Monil(gaster, 48 150, 151, 161, 165, 216, 232, 239, Moniligastridae, 41, 42, 43, 46, 47-8, 66, 240, 241, 251, 255 67, 80, 146, 147 lacteum, 82, 98, 128, 146, 153, 165, 183, Moniligastrinae, 42, 43 228,229,232,241,251,255,256 Monocystls, 19, 148 tyrtaeum, 62, 256 Monogaster, 53 oculala, Eophtla, 256 morrisl, Pheretima, 63, 80, 247 oculatus, Helodrtlis, 241, 253, 256 mucronata, Capillaria, 227 Oligochaeta, 22, 40, 41, 42 muldali, Allolobophora, 62, 256 Omodeona, 53 muldali, Bimastos, 241 Onesla sepulchralis, 149 murchiei, HistlOstoma, 148 subalpina, 149 Myxocystis, 148 Onychochaeta, 56 Opisthodrilus, 56 nagana, Drawida, 80 Naididae, 2, 41, 42 palustrls, Blmastos, 62, 249 N annodrilts, 50 paptl/ifer, Drawida, 80 Nellocystis, 148 papllliJer, Plutellus, 61, 79 Nellogaster, 49 Parabursaria, 148 Nelloscolex, 49 Parachilota, 52 Nemantodrilus, 54 paradoxa, Paricterotaenia, 226 NematocystlS, 148 Parascolex, 55 Nematoda, 226 parasltlca, Frtdericla, 146 Nematogenia, 50 Parendrilinae, 43,54 Neochaeta, 49 Parendrtlus, 54 N eodrilacae, 44, 52 Partcterotaenia paradoxa, 226 SYSTEMATIC INDEX 307 pan'us, Blmastos, 20, 62, 93, 241, 249, 256 Platydnlus, 54 Paultstus, 51 Platyhelminthes, 148 peguana, Pheretima, 226 platyura, Dendrobaena, 250 Pellogaster, 53 Pleuroij'stlS, 148 pennsyivanica, Fraxmus, 198 Pliconogaster, 50 Penodrtlus, 52 Pliogaster, 50 Perionychini, 45 Plutellus, 41, 52, 61, 65 Perionyx, 7, 49, 95, 114 paptlltfer, 61, 79 excavatus, 80, 95 umbellulartae, 61, 79 mtllardl, 140 Pluvlaits apricarta, 147 Penscolex, 56 Pollenia rud,s, 149 Perzssogaster, 49 Polycercus, 148 Phalans, 164 lumbrict, 226 Pheretlma, 7, 8, 18,21,23,25,30,32,36,37, Polychaeta, 40, 41 38, 39, 47, 49, 66, 67, 73, 78, 81, 88, polychromogenes, Nocardia, 187 100, 101, 107, 110, 137, 148, 184,226, Polytoreutus, 55 229, 245, 247 Pontodnlus, 52 agrestls, 63, 107 bermudenSlS, 61, 79 alexandn, 79 gracilis, 61 anomala, 79 Pontoscolex, 56, 66 blcmta, 63 corethrurus, 61, 66, 79, 228 calyarn/ca, 22, 25, 63, 102, 154, 157, Porrocaecum crassum, 227 158, 247 enslcaudatum, 227 campanulata, 79 posthuma, Pheretlma, 39, 63, 140 (ommuntssima, 93 Priodochaeta, 53 defecta, 79 PrlOdoscolex, 4, 53 dlfjrmgens, 63, 80, 122, 247 Pronaldites, 41 e!ongata, 79, 228 Protoscolex, 41 eXlgus, 79 latus, 41 glabra, 79 Protozoa, 98, 99, 148, 188, 226 hawayana, 63, 80, 247 pudentotectus, Metastrongylus, 226 htlgendorji, 63, 72, 79 pumila, Lennoscolex, 80 houlletl, 63 putorti, Caplllana, 227 hupelensls, 24, 63, 73, 74, 75,127, I31, 138, pygmaea, Dendrobaena, 241, 250 140, 142, 153, 157, 158, 247 Pygmaeodrilus, 5I mfanttllS, 79 Pythium, 188, 224 levIS, 63 lovertdgel, 63 Quechuona, 5 I mendlana, 79 Quedius (Mlcrosaurus) mesomeiinus, 147 minIma, 63, 247 Quercus robur, 198 morrisl, 63, 80, 247 peguana, 226 Ramiella, 53 posthuma, 39, 63, 140 blshambari, 80 rimosa, 79, 80 Ramzellona, 53 rodertCencls, 247 rara, Drawlda, 80 stellert, 8 redii f. gigantes, Hormogaster, 65 vanans, 79 RhabditIs, 148 Virgo, 79 Rhmodrtlus, 56 voeltzkowl, 79 Rhododrilus, 52 zoyslae, 79 rhombeatus, Causus, 147 Pheodrilidae, 42 Rhynocystls, 148 phosphoreus, Mlcroscolex, 61 Rillogaster, 53 Picea manana, 198 nmosa, Pheretlma, 79, 80 pltca, Captllaria, 227 nparta, DIplocardIa, 140, 247 Plaglochaeta, 52 robur, Quercus, 198 Plaglotoma, 148 robusta, Allolobophora, 146 308 BIOLOGY OF EARTHWORMS

roder/eenets, Pherettma, 247 subalpina, Onesra, 149 Rosadrilus, 55 subandina, Eukerria, 79 rosea, Al/olobop'hora, 9, 62, 63, 70, 73, 81, subrubicunda, Dendrobaena, 19, 70, 71, 73, 83, 125, 126, 127, 128, 129, 130, 131, 82, 85, III, 150, 153, 154, 192, 232, 132, 165 240, 256 rosea, Elsema, 62,' 63, 72, 73, 80, 98, 140, subvirrd,s, Trocheta, 147 158, 256 surensis, Oetoehaetona, 80 rostoehiensis, Heterodera, 224 suum, Asmris, 228 rubecula, Erithacus, 147 sylvaticus, Didymogaster, 19 rubel/us, Lumbricus, 62, ~, 70, 71, 73, 80, Sylvodrilus, 52 82, 93, 98, 108, I I I, 127, 128, 145, Syngamus, 148 I~I~I~I~I~I~I~I~ merulae, 227 173, 183, 192, 196, 226, 227, 229, skrjabinomorpha, 227 232, 239, 240, 241, 248 trachea, 225, 227 rublda, Dendrobaena, 62, 80, 82, 127, 131, Syngenodrilinae, 43 145, 146, 149, 150, 165, 17°,232, 239, synoecnema, 148 240, 250, 256 rubida f. subrubicunda, Dendrobaena, 146, Taenia euneata, 148 241, 250, 256 Talpa europaea, 147 rubida f. tenuis, Dendrobaena, 81, 241, 250, Tazelaarra, 5 I 256 Teleudrilus, 55 rudts, Pol/mia, 149 tenuis, Bimastos, 8, 256 termiticola, Notoscolex, 147 salami, Metastrongylus, 226 terrestris, Al/olobophora, 146, 252 saitens, Dichogaster, 63 terrestris f. longa, Al/olobophora, 73 saltensis, Eukerria, 63 trapezoides, Allolobophora, 62, 255 Sareophaga earnaria, 149 f. trapezoldes, caligmosa, Allolobophora, haemorrhoidalis, 149 26, 63, 80, 252, 255 strrata, 149 turgida, A71olobophora, 62, 80, 255 Sehubotztella, 55 f. (ypica, caliginosa, Allolobophora, 63, dunguensts, 78 252, 255 Seolail/us, 54 terrestns, Lumbricus, 6, 13, 21, 27, 39, 62, Seolioscolides, 53 72, 76, 80, 81, 82, 83, 84, 85, 86, 88, scutulum, Testacella, 147 89, 90, 91, 92, 93, 94, 97, 98, 103, sepulchralis, Onesia, 149 106, 107, 108, I II, 114, 115, 117, 125, Serratia'marcessens, 184 126, 128, 129, 130, 131, 132, 134, singularis, Diplocardia, 61, 247 136, 138, 139, 140, 145, 146, 148, 149, skrJabmomorpha, Syngamus, 227 150, 151, 152, 153, 154, 156, 157, 160, smlthii, Sparganophilus, 63 161, 162, 166, 168, I~, 172, 173, 177, sonomae, Sparganophllus, 63 178, 179, 182, 183, 184, 185, 188, 192, Sparganophilidae, 47,57, 63, 65, 245, 248 195, 196, 198, 199, 200, 201, 206, 216, Sparganophilinae, 43 218, 21 9,226,227,228,229,230,231, Sparganophilus, 57, 245, 246 232, 234, 235, 236, 237, 239, 240, 24 1, elSent, 63, 154, 248 242, 248 smithii, 63 Testacella hallOlidea, 147 sonomae, 63 mauger, 147 S penceriel/a, 53 scutulum, 147 S phaeractinomyzon, 148 tetraedra, Etseniella, 62, 80, 165, 232, 240, Spirochaeta, 148 241, 249 Spiroptera, 148 f. hercynia, Eiseniella, 241, 249 turdi, 227 f. typrea, Elseniel/a, 165, 241, 249 stellerr, Pheretlma, 8 Thamnodrrloldes, 56 Stephanurus, 148, 227 Thamnodrilus, 56 striata, Sarcophaga, 149 Thatonta, 51 Streptomyces coellcolor, 187 Teleuloreutus, 55 Sturmus vulgaris, 147 Thelohanta, 148 SYSTEMATIC INDEX 309

Thommx, aerophtlus, 227 veneta f. typtca, Dendrobaena, 241, 250, 256 Til/etla controversa, 188, 224 reneta var. hortensts, Dendrobaena, 241 Tonoscolex, 50, 53 veneta var. htberntca f. dendrotdea, Dendro- trachea, Syngamus, 225, 227 baena, 241 tncolor, Hystrichus, 226 veneta var. htbernica f. typlca, Dendrobaena, Trtgaster, 53 146, 241 Trocheta subvtrldts, 147 veneta var. zebra, Dendrobaena, 241 tuberculata, Allolobophora, 62, 80, 93, 241, Ventura inaequalis, 188 252 Virgo, Pherettma, 79 Tubificidae, 42 voeltzkowi, Pheretima, 79 tumtdis, Bimastos, 62, 82, 85, 249 vulgaris, Sturmus, 147 turdi, Sptroptera, 227 Turdus ericetorum, 147 waltoni, Eutyphoeus, 78, 142 Turdus merula, 147 Wegenerzella, 53 turgida, Allolobophora, 62, 80, 255 Wegenenona, 53 tyrtaeum, Enterion, 256 welcht, Bimastos, 62 tyrtaeum. OctolastOn, 62, 256 Woodwardtella, 50

Yagansta, 52 Udema, 52 avesicula, 79 Zapotecta, 52 undula, Dilepus, 226 Zea mays, 228 Uropoda agitans, 149 zebra, Eisenta, 62 zeteki, Bimastos, 62, 72, 74, 75, 85, 86, 88, vartans, Pheretima, 79 89, 145,249 veneta, Eisenza, 226, 256 zoyslae, Pheretima, 79 Author index

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

Abrahamsen, G., 123, 146, 261 Bather, E. A., 41, 263 Agarwal, G. W., 228, 261 Bauer, K., 212, 213, 263 Aichberger R. von., 184, 261 Baweja, K. D., 188, 224, 263 Aisyazhnyuk, A. A., 261 Baydonabiciene, Z., 262 Aldag, R., 196, 261 Baylis, H. A., 142, 146, 263 Alicata, J. E., 294 Beauge, A., 144, 263 Allee, W. c., 149, 261 Bejsovec, J., 228, 264 Allen, R. W., 261 Beiousova, N. K., 124, 26Q Almeida, A. M., 97 Bengston, S.-A., 147, 264 Anderson, H. L., 282 Benham, W. B., 13, 29, 41, 264 Anstett, M., 261 Beynon, K., 270 Arbit, ]., 93, 261 Bharucha-Reid, R. P., 93, 264 Aristotle, xv Bhatt,]. V., 174, 184, 188,224,280 Arldt, T., 41, 261 Bigger, J. H., 213, 264 Arlidge, G. Z., 214, 280 B1ancke, E., 175, 190, 264 Arnold, M. K., 214, 269, 276 Blankwaardt, H. F. H., 212, 264 Arrhenius, 0., 149, 261 Block, W., 123, 264 Arthur, D. R., 99, Il3, 209, 210, 241, 262 Bocock, K. L., 264 Arthur, J. H., 271 Bodenheimer, F. S., 149, 194, 264 Ash, J. S., 266 Boice, R., 94, 140, 292 Aspock, H., 215, 281 Bollen, W. B., 181, 289 Atlavinyte, 0., xvii, 174, 183, 197, 230, Bornebusch, C. H., 118, 124, 149, 264 262 Bouche, M. B., xvii, 122, 264 Avei, M., 262 Boyd, ]. M., 163, 264, 265 Axeisson, B., 209, 262 Boykins, E. A., 217 Boynton, D., 159, 265 Bacheiier, G., xvii, 149, 163, 210 Bradley, ]., 192, 196, 266 Bahl, K. N., 18, 73, 102, 140, 262 Brain, R. c., 273 Baker, W. L., 262 Braunig, G., 98, 107, 284 Bakhtin, P. U., 192, 193, 263 Bray, J. R., 173, 265 Baldwin, F. M., 90, 94, 263 Bretnall, G. H., 265 Ball, D. F., 289 Bretscher, K., 118, 265 Ball, R. c., 263 Breza, M., 226, 265 Baluev, V. K., 196, 263 Brinkhurst, R. 0., 61, 265 Banage, W. B., 123, 264 Brown, B. R., 166, 265 Barker, R. ]., 213, 215, 263 Brown, D. M., 265 Barley, K. P., 118, 124, 138, 164, 165, 167, Bruel, W. E. van der, 187, 265 172, 173, 174, 176, 179, 189, 197, 205, Briisewitz, G., 184, 185, 186, 265 206, 228, 229, 263 Buahin, G. K., 211, 265 Barrett, T. j., xvii, 199, 263 Budaviciene, I., 262 Bassalik, K., 182, 184, 190, 192, 263 Buntley, C. J., 265 AUTHOR INDEX 3 I I

Byzova, Y. 8., 97, 98, 213, 265, 274 Douhalei, N., 281 Dowdy, W. W., 131, 157, 269 Cain, A. J., 265 Dreidax, L., 144, 196, 269 Capstick, C. K., 264 Drift, ]. van der, xvii, 212, 214, 264, 268, Carter, G. 5., 1I6, 266 269, 291 Causey, D., 226 Dubash, P. J., xvi, 77, 78, I16, 296 Cernosvitov, L., 53, 60, 146, 255, 266 Duley, F. L., 296 Chadwick, L. c., 192, 196, 266 Dustman, E. H., 215, 269 Chandler, R. F., 120, 269 Dutt, A. K., 192, 193, 201, 269 Chapman, G., 266 Dzangaliev, A. D., 124, 269 Chen, C. M., 222, 266 Christensen, R. E., 274 Eaton, T. H. Jr., 120, 269 Cialdini, R. 8., 290 Eberhardt, A. 149, 269 Clebsch, E. E. G., 290 I., Cockerell, T. D. A., 149, 266 Edwards, C. A., 1I9, 120, 124, 151, 178, Cohen, S., 103, 266 179, 199,2°3,206,2°9, 21I, 212, 213, Coin, C. ]., 266 214,215,217,218,222,237,265,267, Combault, A., 26, 196, 266, 291 269, 270, 276 Ehlers, W., 206, 270 Compton, O. c., 159, 265 EI-Duweini, A. K., 22, 102, 118, 125, 150, Conder, P. J., 215, 266 Cori, C. J., 227 152, 154, 156, 157, 158, 161, 270 Ellenby, 224, 270 Cossens, G. G., 194, 197, 201, 295 c., Ellis, D. ]., 283 Couperus, H., 97, 268 Empson, D. W., 269 Couture, G., 219, 283 Cragg,]. B., 266 Escherich, K., 147, 271 Cramp,S., 215, 266 Escrit, J. R., 209, 210, 211, 271 Crompton, E., 210, 267 Evans, A. c., xvii, 60, 62, 68, 69, 71, 72, Crossley, D. A., 166, 173,219,267, 270 81, 82, 84, 85, 87, 120, 125, 131, 132, Curry, L. L., 263 133, 134, 136, 139, 140, 141, 144, 145, Czerwinski, Z., r81, 267 1~lnl~I~I~I~I~I~ 172, 190, 194,203,205,229,231,232, 1 DaCiulyte, Y. A., 230, 262 255, 266,27 Darwin, c., xii, xvii, 90, 108, 144, 165, 228, 267 Feldkamp. ]., 5, 27' Datta, L. G., 94, 267 Fenton, G. R., 271 Davey, S. P., 212, 267 Finck, A., 151, 181, 192, 271 Davies, H., 267 Fleming, W. E., 213, 271, 272 Davis, 8. N. K., 215, 217, 267 Ford, J., 1I8, 272 Dawson, A. 8., 35, 267 Fox, C. ]. S., 212, 272 Dawson, R. c., 184, 192, 193, 267 Franz, H., 173, 272 Day, G. M., 174, 185, 267 Fraser, C. H. T., 93, 272 Decker, G. c., 213, 217, 264, 283 French, M. c., 217, 267 Dennis, E. 8., 212, 213, 269 Fujita, 0., 93, 278 Devigne, J., 267 Dhawan, C. L., 267 Dhennin, L., 225, 267 Gaddie, R. E. Sr., xvii, 199, 272 Dingwall, A. R., 128, 197,276 Gansen, P. S. van, 99, 100, 272 Dixon, R. M., 194, 288 Garner, M. R., 139, 239, 272 Djuffill, R. A. Jr., 297 Gast, J., 165, 272 Doane, C. c., 213, 215 Gates, G. E., xiii, 41, 43, 44, 46, 50, 51, Dobson, R. M., 199, 268 52, 57, 58, 61, 62, 63, 64, 65, 66, 79, Doeksen, J., xvii, 84, 95, 97, 119, 151,268, 80, 81, 82, 85, 95, 115, 136, 142, 255, 29 1 256, 272, 273 Doerell, E. c., 210, 268 Gavrilov, K., 62, 63, 79, 273, 279 Dotterweich, H., 151, 268 Genov, T., 226, 273 Douglas, D. E., xvii, 199, 272 Geoghegan, M. ]., 193, 273 312 BIOLOGY OF EARTHWORMS

Gerard, B. M., xvii, 68, 72, 84, 126, 129, Hess, W. N., 31, 33, 35, 107, 276 130, 133, 135, 136, 152, 153, 155,255, Heungens, A., 170, 213, 214, 215, 277 273, 274 Hill, ]. P., 148, 276 Gersch, M., 230, 274 Hirst, ]. M., 188, 277 Gewehr, H., 293 Hobmaier, A., 226, 277 Ghabbour, S. I., 102, 118, 125, 144, 150, Hoeksema, K. ]., 192, 277 152, 154, 156, 157, 158, 161, 187,27°, Hoffman, ]. A., 188, 224, 277 274 Hogben, L., 157, 277 Ghilarov, M. S., xvii, 184, 195, 200, 213, Hogg, T. W., 230 274 Hollister, P. L., 261 Ghoca, M. L., 290 Hook, R. J. van, 230, 277 Giesecke, F., 175, 190, 264 Hopkins, A. R., 213, 214, 277 Gilbert, 0., 264 Hopkins, H. T., 192, 203, 204, 206, 207, Gish, CD., 216, 217, 218, 220, 274 212, 277 Gissel-Neilson, G., 287 Hopp, H., !I 8, 133, 136, 156, 192, 194, Glasgow, L. L., 218, 295 196, 199, 203, 204, 206, 207, 208, 277, Goffart, H., 213, 214, 274 295 Gordon, W. T., 278 Howell, CD., 94, 107, 277, 278 Gorham, E., 173, 265 Hoy, H. M., 213, 278 Graff, 0., xvii, 72, 155, 157, 176, 181, 196, Hubl, H., 106, 277 203, 205, 239, 25 1, 274, 275 Huhta, V., 209, 278 Grant, W. C, 24, 74, 127, 131, 138, 140, Hunt, L. B., 215, 217, 278 152, 153, 157, 181, 275 Hutchinson, S. A., 185, 224, 278 Grassi, B., 226, 275 Hyche, L. L., 214, 278 Gray,]., 113, 114, 275 Hyman, L. H., 85, 278 Greenwood, D. E., 213, 275 Griffiths, D. C, 214, 275 Inoue, T., 278 Grigor'eva, T. G., 213, 275 Ireland, M. P., 220, 221, 278 Grove, A. ]., xvii, 15, 16, 18, 28, 29, 32, Isa, A. L., 282 37, 275 Iwahara, S., 93, 278 Guild, W. J., xvii, 62, 68, 69, 71, 72, 81, 82, 84, 87, 120, 125, 127, 131, 132, Jacks, G. V., 94, 278 133, 134, 136, 141, 144, 145, 150, 153, Jacob, A., 209, 210, 279 154, 155, 162, 164, 165, 172, 173, 192, Jacobson, G. M., 176, 180, 181, 201, 283 194, 203, 205, 229, 231, 232, 27 1, 275 Jakubczyk, H., 267 Gunthart, E., 2I)', 276 Jamieson, B. G. M., 45, 46, 61, 265, 279 Gurianova, O. Z., 192, 276 Janda, V., 62, 279 Jeanson-Luusinang, C, 150, 185, 279 Hadley, C H., 213, 271 Jefferson, P., 120, 205, 210, 279 Hamblyn, C J., 127, 128, 197, 276 Jennings, A. C, 176, 189, 263 Handa, B. K., 267 Jevniaux, C, 267 Handley, W. R. C, 271 Joachim, A. W. R., 192, 279 Hanel, E., 90, 276 Johnson, M. L., 98, 279 Hanley, I. M., 265, 272 Johnstone-Wallace, D. B., 119, 208, 279 Hardman, ]. A., 154, 215, 216, 217, 218, Jongerious, A., 277 298 Joshi, N. V., 190, 196, 279 Harman, W. ]., 276 Julin, E., 64, 279 Harmsen, G., 189, 276 Harrison, R. B., 215, 267 Kahsnitz, H. G., 196, 279 Hasenbein, G., 230, 276 Kalmus, H., 279 Haswell, W. A., 148, 276 Kamel, M., 185, 188, 224, 278 Heath, G. W., 124, 166, 168, 169, 170, 176, Kanwar, ]. S., 181, '90, 192, 287 209, 237, 269, 270, 276, 280 Karmanova, E. M., 226, 227, 279 Heck, L. von, 92, 93, 276 Karpachevskii, L. 0., 288 Heimburger, H. V., 276 Keilin, D. 280 Hensen, V., xvi, 276 Kelkar, B. V., 190, 196, 279 AUTHOR INDEX 313

Kelly, W. A., 291 Luckman, W. H., 213, 217, 283 Kelsey, j. M., 214, 280 Lugauskas, A., 183, 262 Kevan, D. K. McE., xvii, 280 Lukose, J., 28 3 Khambata, S. R., 174, 184, 188,224, 280 Lund, E. E., :zz6, 283 King, H. G. c., 166, 169, 276, 280 Lunt, H. A., 176, ISo, 181, 201, 283 Kirberger, c., 97, 280 Lyons, C. H., 214, 296 Kirk, R. L., 157, 214, 277 Kirk, V. M., 277 Madge, D. 5., xvii, 75, 83, 123, 142, 144, Kleinig, C. R., 172, 179, 197, 263, 280, 287 ISO, 154, 157, 173,283 Kleist, S. M., 290 Magalhaes, P. 5., 226, 283 Knop, J., 29, 280 Maldague, M., 219, 283 Kobatake, M., 187,280 Mamajev, B. M., 200, 274 Kobayashi,S., 63, 280 Mamytov, A., 192, 193, 283 Kollmannsperger, F., 133, 144, ISS, 163, Mangold, 0., 108, 165, 166, 283 280 Marapao, B. P., 106, 283 Kollmannsperger, G., 144, 280 Marshall, V. G., 198, 283 Kondo, K., 222, 278 Martin, A. W., 102, 283 Korschelt, E., 72, 280 Martin, ]. P., 193, 297 Kozlovskaya, L. 5., 183, 185, 280 Matlu, N., 278 Kring, ]. B., 214, 215, 281 Mayne, D. W., 295 Krivaneck, J. 0., 93, 280 McCalla, T. M., 205, 296 Kriiger, F., 96, 124, 281 McInnes, D. c., 297 Kubiena, W. L., 163, 281 McInroy, D., 229, 284 Kiihnelt, W., xvii, 163, 281 McLeod,]. H., 147, 284 Kurcheva, G. F., 169, 281 Meer, K. van der, 277 Meggitt, F. J., 226, 284 Lan, H. van der, 281 Mellanby, K., 284 Ladell, W. R. 5., 1I9, 281 Mendes, E. G., 97, 284 Lahr, ]. P., 261 Merker, E., 98, 107, 284 Lakhani, K. H., 121, 177, 281 Meyer, L., 190, 284 Lal, R., 206, 281 Michaelsen, W., 40, 41, 42, 47, 61, 64, Lauer, A. R., 93, 281 284 Lavelle, P., 74, 136, 138, 144, 153, 281 Michon, ]., 72, 73, 74, 84, 86, 153, 155, Laverack, M. S., xvii, 17,96, !O3, !O8, 166, 284 281, 282 Miles, H. B., 188, 284, 285 Lawrence, R. D., 175, 282 Millar, H. R., xvi, 175, 281 Lee, K. E., xvii, 43, 46, 84, 195, 229, 282 Millott, N., 99, 285 Legg, D. c., 213, 215, 282 Minderman, G., 268 Leitenberger, L., 173, 272 Moment, G. B., 85, lIS, 285 Lesser, E. j., 98, 282 Monnig, H. 0., 226, 285 Lewis, H. B., 103, 175, 266 Moore, A. R. 285 Lidgate, H. ]., 213, 282 Moore, B., 109, 149, 285 Lindquist, B., 165, 175, 282 Morris, H. M., 119, 205, 285 Lipa, J. ]., 213, 282 Morrison, F. 0., 213, 285 Lissman, H. W., 113, 114, 275 Mozgovoy, A. A., 227, 285 Liv, C. L., 222, 266 Muldal, 5., 62, 79, 285 Ljungstrom, P.O., xvii, 61, 66, 79, 142, Miiller, G., xvii, 285 147, 229, 282 Miiller, P. E., 165, 195, 285 Lofty, J. R., 120, 124, 151, 156, 160, 161, Murchie, W. R., xvii, 61, 62, 71, 75, 84, 199,203,205,206,209, 21I, 212, 214, 85, 86, 89, 127, 136, 145, 285, 286 219, 268, 269, 270, 275, 282, 290 Lohm, u., 262 Nakamura, Y., 137, 138, 149, 286, 296 Long, W. H., 213, 282 Nathans, 5., 118, 124, 291 Love, C. W., 265 Needham, A. E., 103, 173, 175, 176, 177, Low, A. ]., 118, 192,283 286 Lowe, D. G., 293 Negi, L. 5., 261 314 BIOLOGY OF EARTHWORMS

Nelson, J. M., 1I8, 286 Rao, K. S. K., 261 Newell, G. E., xvii, 15, 16, 18, 28, 29, 32, Raw, F., 1I9, 120, 122, 124, 125, 159, 168, 37, 1I2, 275, 286 169, 170, 173,21 1,214,221,275,289, Nicol, H., 210, 287 290 Nielson, C. 0., 196, 197, 199, 220, 287 Reichle, D. E., 267, 270 Nielson, M. G., 287 Reinecke, A. ]., 142, 147, 157, 229, 282, Nielson, R. L., 286 290 Nijhawan, S. D., 181, 190, 192, 287 Ressler, R. H., 94, 290 Nilsson, A., 264 Reynolds, J. W., xvii, 62, 63, 79, 122, 124, Noble, J. c., 197, 202, 287 147, 229, 290 Nordstrom, S., 146, 156, 161,264, 287 Reynolds, W. M., 229, 290 Nowak, E., 267 Reynoldson, T. B., 1I8, 124, 290, 291 Rhee, J. A. van, xvii, 118,' 124, 126, 173, O'Connor, F. B., 291 196, 198, 200, 221, 290 Ogg, W. G., 210, 287 Rhoades, W. c., 214, 291 Ogles, G. D., 149, 296 Ribaudcourt, E., 196, 291 Oldham, c., 287 Richards, ]. G., 197, 291 Oliver, J. H., 148, 287 Richardson, H. c., 150, 159,210,211,291 Olson, H. W., 149, 152, 287 Richter, G., 213, 291 Omodeo, P., 43, 44, 46, 62, 65, 255, 287 Robertson, J. D., 99, 29 1 Otanes, F. G., 222, 287 Robinson, J. S., 92, 93, 292 Rodale, R., xvii, 210, 213, 292 Panditesekera, D. G., 192, 279 Rodionova, L. Z., 288 Papedick, R. I., 265 Roots, B. I., 100, II 0, 152, 292 Parker, G. H., 1I0, 288 Rossenkoetter, ]. S., 94, 140, 292 Parle, J. N., 182, 183, 186, 187, 193, 288 Rovelli, G., 226, 275 Parshley, H. M., 110, 288 Roy, S. K., 141, 144, 292 Patel, H. K., 213, 222, 288 Rundgren, S., 131, 146, 156, 161, 264, Pathak, A. N., 190, 294 287, 292 Peachey, J. E., 288 Ruschmann, G., 175, 187, 292 Pensson, T., 262 Russell, E. J., xii, xvi, 159, 175, 196, 292 Peredel'sky, A. A., 218, 288 Rysav)', B., 226, 227, 292 Perel, T. S., 170, 288 Ryzhikov, K. M., 227, 292 Peterson, A. E., 195, 288 Petrov, B. c., 149, 288 Sacho, R. ]., 217, 278 Phillips, E. F., 149, 288 Salisbury, E., xvi, 149, 151, 200, 292 Pickard, J. A., 213, 215, 217, 296 Saroja, K., 292 Piearce, T. G., 146, 151, 165, 288 Sastry, K. S. S., 222, 228, 289 Polivka, J. B., 21 I, 213, 214, 289 Satchell, J. E., xi, xvii, 70, 72, 81, 84, 85, Polsky, M. N., 192, 193, 263 86, 108, 109, 1I8, 120, 121, 123, 127, Pomerat, G. M., 97, 289 128,129,149,150,151,155, 159,161, Ponomareva, S. I., 181, 182, 192, 193, 206, 163, 165, 166, 16g, 173, 175, 177, 178, 289 179, 180, 182, 188,205,275,281,286, Poryadkova, N. A., 288 293 Powers, W. L., 181, 289 Saussey, M., 146, 195, 293 Prabhoo, N. R., 289 Scharpenseal, H. W., 293 Pratt, K. c., 85, 296 Schmid, L. A., 106, 293 Prosser, C. L., 289 Schmidt, H., 92, 294 Puh, P. c., 151, 181, 289 Schneider, K. c., 34, 294 Purdy, L. H., 188, 224, 277 Schread, J. c., 142, 213, 214, 223, 294 Puttarudriah, M., 222, 228, 289 Schrevan, D. van, 189, 276 Schwartz, B., 226, 243, 294 Raffy, A., 97, 289 Scopes, N. E. A., 214, 298 Ragg, J. M., 289 Scott, H. E., 214, 294 Ralph, C. L., 97, 289 Scrickhande, J., 190, 294 Ramsay, J. A., 101, 102, 110, I I I, 289 Sharma, R. L., 267 AUTHOR INDEX 315

Shindo, B., 294 Urquhart, A. T., xvi, 297 Shiraishi, K, 98, 159, 294 Sims, R. W., xvii, 43, 44, 45, 46, 294 Vail, V. A., 82, 85, 89, 297 Singh, A., 267 Villot, F. C. A., 226, 297 Sison, P. L., 222, 288 Vimmerstedt, l P., 202, 297 Skarbilovic, T. S., 227, 294 Vogel, R., 226, 297 Slater, C. S., 194, 196, 199, 204, 210, 277, Volz, P., 195, 297 294, 295 Smallwood, W. M., 33, 105, 295 Waid, J. S., 264 Smith, F., 295 Waite, R. H., 297 Smith, R. D., 218, 295 Waksman, S. A., 189, 193, 296, 297 Sokolov, D. F., 288 Walton, W. R., 119, 222, 298 Stafford, C. J., 270 Ward, W. c., 277 Stephenson, J., xvi, xvii, 10, 12, 13, 16, 18, Watanabe, H., 73, 298 26, 27, 36, 38, 40, 41, 42, 43, 44, 47, Waters, R. A. S., 133, 138, 139, 165, 197, 64, 65, 81, 89, 98, IIO, II4, 148,229, 198, 205, 298 257, 295 Watkin, B. R., 208, 298 Stickel, L. F., 215, 269, 295 Way, M. ]., 214, 298 Stickel, W. H., 295 Weber, G., 213, 214, 298 Stock dill, S. M. J., 195, 197, 201, 295 Wehr, E. E., 283 Stockli, A., II8, 144, 175, 181, 182, 186, Weisbach, W. W., 229, 298 191, 194, 295 Went, J. c., 184, 185, 298 Stokes, B. M., 295 Westeringh, W. van def, 2 II, 298 Stolte, H. A., xvi, 148, 228, 295 Wheatley, G. A., 215, 216, 217, 218, 298 Stone, P. c., 149, 296 Wherry, E. T., 149, IS0, 298 Stop-Bowitz, c., 296 White, G., xi Storey, 1. F., 277 Whiting, A. E., 270 Stough, H. B., 34, 296 Whitney, W. K, 214, 298 Stringer, A., 2 I 3, 214, 2 I 5, 217, 296 Wiegland, K., 209, 210, 279 Sun, K. H., 85, 296 Wilcke, D. E. von, 72, 73, 118, 298 Svendsen, l A., II8, 122, 124, 127, 165, Wilkinson, G. E., 195, 206, 298 296 Wilcox, H. G., 277 Swaby, R. J., 192, 296 Wingerden, C. G. van, 84, 151, 268 Swartz, R. D., 93, 296 Witkamp, M., 177, 299 Wittich, W., 108, 166, 177, 299 Takano, S., 149, 296 Wojewodin, A. W., 212, 299 Tembe, V. B., xvi, 77, 78, 117,296 Wolf, A. V., 110, 156,299 Tenney, F. G., 189, 296 Wollny, E., 194, 196, 299 Teotia, S. P., 183, 190, 192, 193, 194, Woodhead, A. A., 227, 299 205, 296 Woodman, M. G., 264 Thomson, A. J., 140, 297 Wooton, R. l, 220, 278 Thompson, A. R., 213, 214, 215, 217, 270, Wright, M. A., 166,299 296 Tillinghast, E. K., 102, 103, 297 Yegorova, S. V., 288 Tischler, W., 205, 297 Yerkes, R. M., 90, 92, 93, 299 Torvic, M. M., 261 Tracey, M. V., 99, 297 Zajonc, 1., xvii, 124, 209, 299 Trifonov, D., 222, 297 Zarrow, M. T., 97, 289 Tromba, F. G., 227, 297 Zhdannikova, E. N., 183, 185, 280 Twinn, D. c., 264 Zhinkin, L., II6, 299 Zicsi, A., xvii, 118, 122, 204, 222, 299 Uhlen, G., 210, 297 Zrazhevski, A.!., 183, 184, 198, 299 General index

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

Absorption zone, 100 Algeria, 58 Acclimation, 157 Alimentary canal, I, 20-2, 23, 34, 37, 100, Accessory reproductive bodies, 8 148, 183, 184, 185 Acetycholine, 106, 116 innervation of, 33, 34 Acid intolerant species, 150 plexus, 25 tolerant species, 149, 150 Alkaline soils, 234 soils, 149, 150, 151, 197, 210, 234 Alkaloid substances, 108 reaction of worms to, 127, 243 Allantoin, 100, 175 Acids, 98, 108, 151, 175 Alluvium, population in, 162, 163 Actinomycetes, 182, 183, 185, 186 America, 55, 57, 74 Activity, 138, 145, 146, 154, 178 Central, 5', 52, 56, 57, 65, 66, 67 cycles, 93, 97, 106 North, xv, 52, 53, 59, 61, 64, 144, 205, effects of temperature on, 133, 155, 156 246, 247, 248 seasonal changes in, '3', 132, 134, 136 South, 50, 51, 52, 56, 57, 59,65,66,67, Adders, 147 246, 248 Adolescence, 87 Ammonia, 100, 101, 102, 103, 152, 175, Adrenaline, 106 176, 186 Adverse conditions, 70, 83, 225 Ammonium, 176, 189 effects on soil, 228-9 sulphate, 172, 211 periods, 83 Amoebocytes, 19, 101 Aeration, 127, [59, [94-5, 205, 228 Amphimitic reproduction, 79 Aestivation, 83-4, 152 Ampulla, 29 Afferent nephridial vessel, 18 Amputation, 80 typhlosolar vessel, 18 Amylase, 99, 142 Africa, xv, 49, 50, 53, 54, 55, 56, 57, 58, Anabiosis, 110 59, 60, 65, 66, 74, 125, 136, 142, 147, Anaerobic conditions, 95, 116 152, 222, 229, 246 Anaerobic respiration, 98 Age-class composition, u6 Anaesthesis, 113 African night crawler, 66 Andes, 56 Aggregates, 187, 190, 193, 194, 198, 201 Antarctica, 67 formation of, [92-4 Antibacterial extracts, 187 Agriculture, effects on earthworms, 128, Antibiotic, 187 '45, 203-2[ Anus, I, 20, 85, 95, 100, IIO, 248 Agricultural practice, 47 Apertures, male and female, 9, 12 soils, 152, 228 spermathecal, 12 Albumen cells, 14, 15 Apple scab, 188, 2 I 5 Alcohol, as a preservative, 233 Aquatic species, 9, I I, 43 Alder, 173 Arable, 119, 122, 124, 133, 144, 145, 146, Aldicarb, 2 I 5 156, 160, 161, 193,203,205,216,222, Aldrin, 212, 213, 214, 218 233, 235 Alfalfa, 192, 206 Arciform muscles, 76 GENERAL INDEX 317

Arctic forest, 173 Blackhead, 226 Argentina, 51, 55, 56 Bladder, 102, 103 Artesian irrigation, 206 Blood, 100, 101, 104, 110 Arthropods, 174, 182 plasma, 96 Ascospores, 188 supply, subcutaneous, 27 Asexual reproduction, 79, 80, 8 I system, 103 Ash, 166, 177, 237 vessels, 14, 15, 18, 22, 26, 33, 116 Asia, 49, 56, 59, 64, 67, 246 Blue-green algae, 184 Asia Minor, 59, 64 Body cavity, 9 Association neurones, 105 fluids, 97, I I I Atrazine, 2 I 2 length of, 60 Auckland,s I number of segments of, e5 Australasia, 65, 67 temperature of, 153 Australia, 49, 50, 51, 53, 59, 60, 67, 124, weight of, 110, 152, 173, 174 138, 159, 179, 202, 206 Body wall, 4, 14-17, 15, 19, 22, 33, 34, 99, Austria, 58 101, 103, 105, 110 Autotomy, 85, 89 nerves supplying, 105 Available mineral nutrients, 180-1 Bolivia, 56 A voiding reaction, 151 Borneo, 48, 66 Axon, 34 Bowling greens, 223 Azinphosmethyl, 94, 214 Brazil, 50, 51, 52, 53, 56, 57 Azores, 59, 60 Breeding, 73, 199, 200 Breeding period, 12, 73, 125 Bacteria, 19, 29, 98, 99, 148, 166, 174, 182, Bristle worms, 40 183, 185, 186, 187, 188, 193 Brown bodies, 19 Bacterial gums, 194 Brown earth, 123 Bacteriocide, 187 Bronchial dilation substance, 229 Bacteroidal cells, 101 Buccal cavity, 17, 20, 23, 24, 33 Badgers, 147 innervation of, 35 Bahamas, 48, 50 Buccal chamber, 98, 108 Bahlia, 52 epithelium, 17 Bait, earthworms as, 229 Bulgaria, 222 Bardsey Island, 124 Bullock droppings, as food, 72, 164, 231 Barley, 196, 197 Burma, 48, 49, 50, 57, 95, 142 Basal cells, 14 Burrowing, 110, 138-45, 171, 172, 183, membrane, 17, 33 190, 194 Basalt, 190 experiments, 237-9 Base exchange capacity, 180, 20 I speed of, 139 Basement membrane, 16 Burrows, 88, 89, 90, 91, I II, 119, 121, 122, Bassia tree, 120 128, 139, 191,211,216,222,235,237, Beans, 228, 237 243 Beech, 124, 166, 169, 170, 237 depth of, 140 Beet, 166, 237 diameter of, 140 Behaviour, 88-95, 93, 94, 110 permanent, 129 Behavioural experiments, 88, 243-4 Bursa propulsoria, 79 Benomyl, 214 Butyric acid, 183 Benzopyrine, 230 Bermuda, 56 Caecae, of intestine, 22 BHC, 213 Caecum, 24 Biocides, 2 I I Caesium, 219 Biology xvii, 68-95 Cage for studying earthworm activity, 238 Biomass, xv, 123, 179 Cages, 169 Birch, 166, 170, 177, 237 Calcareous sand, populations in, 163 Birds, 147, 215, 220, 223, 225, 226, 227 Calciferous gland, 20, 22, 44, 45, 46, 50, Blackbirds, 147 55, 57, 58, 59,99, 146, 151, 193,246 Black Spruce, 198 tissues, 20 318 BIOLOGY OF EARTHWORMS

Calcium, 99, 146, 165, 180, 220, 221 drainpipe, 29 carbonate, 19, 99 epithelial, 14, 27 humate, 193 epidermal, 105 metabolism, 146 epithelial brush border, 99 California, Lower, 50, 51, 52, 65 excretory, 100 Cambridge, 124 fibrous, 34 Cameroons, 53, 54, 55, 57, 144 ganglia, 34 Campbell Islands, 5 I giant, 35 Canada, 64, 140 gland, 14, 22, 38, 39, 79, 175 Canary Islands, 59 goblet, 14, 15, 39, 99 Cape Verde Islands, 5 I marginal, 27, 28 Capillaries, 25, 26, 100, 104 mucous, 14, 38, 39 intra-epidermal, 26 nerve, 34, 106 Capillary blood vessels, 14 peritoneal, 19, 101 Capillary water, 194 photoreceptor, 15, 16, 107 Carabid beetles, 147 proprioceptor, 35 Carbamate, fungicides, 2 I 4 reception, 114 insecticides, 2 I 5 replacing, 14 Carbaryl, 215, 223 reproductive, 75 Carbofuran, 214, 215 secretory, 106 Carbohydrates, soluble, 166 sensory, 14, IS, 16, 17, 34, 106 Carbon, combustion, 178, 179 sperm, 39 Carbon nitrogen ratio, 177-80, 181, 202 supporting, 14, 16 dioxide, 96, 98, 159, 178 uric, 101 Carnivorous slugs, 147 Cellulase, 99, 144 Caroline Islands, 48 Cellulose, 177, 182 Carrots, 228 Centipedes, 147 Casting, 138-45 Central cells, 27, 28 in different habitats, 144 Cephalization, 4, 60 time of day of, 142 Cereal stooks, 222 Castor meal, effect of, 160 Cereals, 193, 206, 207 Casts, 100, 131, 140, 142, 144, 145, 151, Cerebral ganglia, 21, 31, 33, 34, 92, 106, 154, 172, 176, 180, 181, 183, 184, 185, 107, I13, 243 186, 187, 188, 189, 190, 191, 192, 193, Ceylon, xiv, 48, 49, 50, 52, 57, 147 194, 196, 201, 205, 206, 222, 223, 224, Chaetotaxy, 4-7 225, 229, 234 Chemical extractants, 120 microbes in, 184, 186 sampling methods, 120, 121 pH of, 200 stimuli, response to, 89, 122 minerals in, 180, 181 transformation, 102 production of, 234-5 transmitter substances, 106 seasonal production of, 133, 141, 144 Chemoreception, 108-g stability of, 192, 193, 201 Chernozem, 182 Cattle, 173 Chickens, 226, 227 Caucasus, 59, 60, 64 Chile, 51, 59 Cells, albumen, 14, IS, 39 China, 48 central, 27, 28 Chitin, 182 chloragogen, 13, 19, 100, 101, I16 Chitinase, 99, 144 ciliated, 22, 53 Chloragogen cells, 13, 19, 100, 101, I16 clitellar gland, 79 Chloragogenous tissue, 19 cocoon secreting, 39 Chloragosomes, 19, 100 Cells, albumen, 14, 15 Chlordane, 213, 214, 218, 223 basal, 14 Chlorfenvinphos, 214 chloragogen, 13, 19, 100, 101, I16 Chloropicrin, 2 I I ciliated, 22, 53 Chlopropham, 212 coelomic epithelial, 17 Choice experiments, I 13 columnar, 39 Cilia, 27, 29 GENERAL INDEX 319

Ciliated cells, 22, 53 Commissural vessels, 25 tubes, 101, 103 Commissures, 25, 26, 107 protozoa, 148 Comparisons of sampling methods, 122 Circular muscle, 14, 15, 17, 18,22, 26, 31, Compost, 146, 200 32, 34, 36, 38, 112, 114 species, living in, 98, 200 innervation of, 33 Computer techniques, 46 Circulation, 103-4 Concentration factor, 218, 220 Circulatory system, 23, 24, 85, 103 Condensed tannins, 166 Circumoesophageal nerve ring, 114 Conductivity, 119 connectives, 31, 32, 33, 107 Congo, 54, 55, 57 Citric acid, 108 Connective tissue, 17, 34, 38 Clasping, 78 Connectives, circumpharyngeal, 31, 32, Clay, 129, 162, 190 33, 107 chimneys, 142 subpharyngeal, 33 population in, 162, 163 Consumption of soil and litter, 172-4 Clitellar gland cells, 79 Contralateral nerve, 105 segments, 13 Contrasting soils, experiments with, 239 tissue, 39 Control of earthworms, 223 Clitellum, I, 10-13, 36, 38, 39, 48, 49, 55, Copper, 221 59, 60, 73, 76, 78, 81, 84, 86, 245, sulphate, 2 I I 246,247,248,249,250,251,252,253, Copulation,S, 39, 42, 76-83, 77, 82 Copulatory chambers, 81 255 position of, I I, 13 pouches, 78 Clover, 164, 196, 208 setae, 58 Cluster flies, 149 Corpuscular bodies, 19 Cobalt, 168 Corsica, 58, 66 Costa Rica, 50, 53, 57 Cocoon, development time of, 72 Cotton, 217 formation, 10, 78, 79 Creatinine, 101 incubation time, 74 Cretaceous period, 46 production, 10, 68, 69, 70, 73, 77, 106, Crop, 20, 21, 22, 99, 177, 184, 215, 223 155, 231, 240 Crop growing areas, 47 Cocoon production, effects of food on, 72, plants, 109 164 yields, effects of earthworms on, 195-9, effects of soil moisture on, 153 198 effects of temperature on, 68, 83, 155 Cropping, effects of, 194, 206-7, 207, 208 Cocoons, 40, 68, 76, 77, 79, 81, 82, 85, 86, Cross-fertilization, 76, 79 I I I, 119, 128, 138, 148, 154, 155, 164, Crows, 147 23 1,232,24°,242 Crumb structure, 108, 187 seasonal production of, 135, 136 Cuba, 51 secretion of, I Cultivation, effects on earthworms, 203---{" shape, 81 2°4,206 size, 232 Cultivations, 205 wall, 81 Culture, growth in, 85, 86 Coelom, 17-20, 22, 27, 148 methods, 232 Coelomic corpuscles, 29 Cultures, 68, 70, 72, 81, 172, 194,218,219, cavity, I 23 1, 232, 237, 239 epithelial cells, 17 Cutaneous respiration, 20 fluid, 9, 13, 17, 18, 19, 26, 96, 100, 101, Cuticle, I, 14, 15, 16, 20, 22, 26, 27, 38, 96, 110,112,151 99, 110, 175 tubes, 54 Cyanazine, 2 I 2 Coelomocytes, I, 95 C ysticercoid stage, 148 Collagenous fibres, 14 Cysticercus, 225 Colombia, 50, 51, 56, 57 Comensalism, 146 Commercial breeding, 199 D-D,211 320 BIOLOGY OF EARTHWORMS

DOE, 215, 218 Dung, 95, 124, 127, 160, 161, 164, 165, Dead roots, 161 172, 173, 174, 197, 198 Deciduous woodland, 165 pats, 127 Dehydration, 109 Dursban, 214 tropism, 110 Dutch elm disease, 2 I 5 Dendrites, 106 Dwarf bunt, 188, 224 Dendrogram, 46 Dyfonate, 214 Denmark, 124, 149 Depth zones, 128 Earthworms, as bait, I 19 Dermis, 106 as benefactors, 221)-30 Deserts, 6 I, 64, 66 as pests, 222~9 Desmids, 184 control of, 223 Desiccation, 14, 20, 70, 72, 83, 154 Earthworm farms, 229 Development, 73, 74, 86, 116, 239 pies, 229 Diapause, 76, 83~4, 151, 234 to kill, 233 facultative, 70, 83, 84, 153 Ecology, xvii, J J8~67 obligatory, 69, 83, 84, 242 Ecuador, 56 Diatoms, 184 Eelworm cysts, 224 Diazinon, 214 Egg sacs, 54 Dicotyledenous plants, 41 Eggs, I, 40, 224, 228, 242 Dieldrin, 213, 217, 218 dispersal of, 40 Diets, 164, 165 Egypt, 57, 59, 75, 125, 150, 154, 161 Digestion, 22, 98~IOO Elderberry, 166 Dimensions of worms, 241 Electrical inhibition of growth, I I 5 Dimethylbenzanthrene, 230 potential, 107 Dipterous larvae, 148 shock, 93, 243, 244 Direct drilling, effects of, 206 stimuli, response to, 89, J09 Disking, effects of, 203 Electrode, 92, 93, 120, 243, 244 Disease transmission, 223~8 Eleocytes, 19 Dispersive power, 127, 128 Elm, 166, 170, 173, 177, 237 Dissection, 233, 242 Endemic species, 47, 61, 64, 66 Distributions, 48, 49, J27~JI, 149, 159, Endrin, 213, 214, 218, 223 163, 233~4, 242 England, 41, 65, 129, 131, 140, 141, 145, Disulfoton, t16, 214 147 Diurnal activity, 138, 178 Environmental factors, influence of, 70, 72, changes, 110 108, 141)-67 respiratory cycles, 97, 106 Enzymes, 99, 142, 168, 182, 225 weight changes, I II Epidermal cells, 105 Diverticula, of buccal cavity, 20 sense organ, 16 DNOC, 212 Epidermis, 10, 14, 15, 16, 17, 26, 27, 32, Dogs Mercury, 166 34, 36, 39, 96, 106, 110, 175, 252 Dominant species, 145 Epilobous prostomium, 2, 4 Dormancy, 152 Epineurium, 34 Dorsal pores, 8, 9, 10, 19, 20, 26, 55, 57, Epithelial brush border cells, 99 58, 60, 88, 110, 151, 248, 249, 250, cells, 14, 27 25 1, 252, 253, 255 sense organs, 14, 35 of A. rosea, 10 Epithelium, 17, 20, 25, 99 Dorsal vessel, 18,21,22,23,24,25,26,37, Essential intermediate hosts, 225 104 Ethiopia, 55 valves of, 25 Euprostates, 53 Dorso-intestinal vessel, 18, 25, 104 Europe, 47, 58, 59, 60, 64, 65, 144, 149, subneural vessel, 18, 23, 25, 104 181, 19~ 199. 205. 246 Drainage, 194, 205, 228 Excretion, I, 25, Ioo~3, 175. 177 Drainpipe cells, 29 Excretory canals. 30 Droughts, 84, 152 cells, 100 Ducks, 226 ducts. 30 GENERAL INDEX 321

organs, 10, 100, 101 165, 166, 167,205,208,229,231,235, system, 27-]I, 46, II I 239, 242 Experimental cages, 235, 236 adsorption, 22 Experiments with earthworms, 190, 231- earthworms, as, 229 244 effect of supply, 164-7 field, 233-44 preferences, 108 palatability, 236--7 supply, 68, 139, 164-7 physiological, 242 Food material, pot, 197, 213 availability of, 177 standardization of, 94 bullock droppings, 72, 164, 23 I Extraction efficiency, 123 compost, 146 External segmental grooves, 17, 18 dung, IZ7, 160, 164 Extra-oesophageal vessel, 25 fodder, 164 herbage, 127 Facultative diapause, 70, 83, 84, 153 horse droppings, 72, 23 I Faeces, 102, 140, 142, 182, 199, 228 leaf, 89, 90, 98, 127, 235 microbial content of, 182, 186 manure, 93 Fallow, 123, 124, 145, 206 oat straw, 164 Families, 42---{,0 organic matter, 127 Farmyard manure, 70, 205, 239 plant, 89, 90, 98, 108, 164 Fats, 101 roots, 133 Fecundity, effects of food on, 164 sheep droppings, 164, 240 effects of moisture on, 152 straw, 231, 239 effects of temperature on, 154 Foot and mouth disease, 225 Feeding, 82 Forest soil, 123, 193, 194, 201 surface, 89 Formalin, 14, 120, 122, 123, 124,232,233, Female pores, 8, I I, 45, 48, 49, 55, 57, 58, 234, 242 60 sampling, 88, 121, 232 Fenestrae, 34 Formaldehyde, 109 Fertilization, 40, 76--83 Fossil records, 41 internal, 79 Fossil worms, 41 Fertilizer, 151, 198, 199, 207, 209 Fragmentation, 168 Fertilizers, effects of, 207-11 France, 52, 66 mineral, 210 Freshwater worms, 13 ni trogenous, 209 Fried earthworms, 229 organic, 207 Frost, effect of, 156 Fever, 229 Fumigants, 2 I I Fibre, collagenous, 14 Fungal hyphae, 186, 193 intra-epidermal nerve, 105 mycelia, 165 lateral giant, 34, 105 spores, 185, 224 median dorsal longitudinal giant, 105 Fungi, 98, 183, 185, 186, 187, 188, 193, motor, 105 194, 224 nerve, 16, 105, 108 Fungicides, 168 Fibrillae, 34 copper, 211 Field experiments, 233-44 effects on worms, 2II, 214 capacity, 195 studies, 174 Filtration, 102 Gambia, 53, 57 Fish bait, I 19 Ganges valley, 5 I Fistula, 228 Ganglia, 21, 31, 32, 35, 92, 106, 107 Flatworm, 147 cells, 35 Flooded soil, 153 cerebral, 21, 31, 35, 92, 106, 107, 113, Fodder, as food, 164 243 Follicles, 4, 75 sub-oesophageal, 31, 106 Food, 22, 68, 70, 72, 89, 90, 93, 98, 108, sub-pharyngeal, 32 113, 117, 127, 139, 146, 160, 163, 164, ventral, 106 322 BIOLOGY OF EARTHWORMS

Gape-worms, 225 Growth, 85-7 Garlic, 19 effects of food on, 164 Gaseous diffusion, 97 effects of temperature on, 164 Genera, 42---{)O electrical inhibitions of, 115 Generator, 120 increment, 86 Genital and other apertures, 8-10 periods, 71, 72, 73, 155 Genital organs, 36, 40 post emergence, 85 pores, 5, I I Guatemala, 51, 57 seta, 5, 6, 60, 8 I Guinea, 53 systems, 36 Gulls, 147 Geographical distribution, 61-7, 233 Gut, 18, 20, 23, 25, 30, 41, 100, 104, 114, Germany, 58, 65, 124, 144 166, 172, 173, 182, 183, 219 Ghana, 54, 55, 144 contents of, 165, 182, 183, 186, 187 Giant cells, 35 enteronephric, 30 fibres, 34, 35, 86, 105, 106 microflora in, 182 dorsal, 105 micro-organisms in, 183 longitudinal, 105 medial, 105 Habitats, 123, 124, 127, 144, 233 Gizzard, 20, 21, 22, 24, 48, 49, 55, 57, 58, arable land, 233 59, 60, 98, 99, 184 chalk downs, 233 Glaciation, 64 heathland, 144, 233 Gland, calciferous, 20, 22, 44, 45, 46, 55, heterogenity of, 127 57,58,59,99, 146, 151, 193 lakes, 246 cell, 14, 22, 38, 39, 79, 175 moorland, 233 lymph, 24, 101 numbers and weights in, 124 mucous, 26, 96 pastures, 233 papillae, 8 rivers, 246 pharyngeal, 20, 99 streams, 246 prostatic, 46, 48 woodlands, 233 salivary, 99 Haemochromagen, 103 Glandular epidermis, 39 Haemoglobin, 14, 26, 85, 96 secretions, 76 Haemorophilic spp., 65 Glucose, 17, J08, 187 Haemorophobic spp., 65 Glycogen, 97, 101 Hairs, 14 Goblet cells, 14, 15, 39,99 Haiti, 52 Golden plovers, 147 Handsorting, 118-19, 122, 124, 233, 234, Golf courses, 210, 223 242 Gonads, I Hants, 124 Granite particles, 190 Harrowing, 203 Granules, 13 Hawaii, 59 yellow refractive, 100 Hay, 196 Grape husks, 189 Hearts, 23, 24, 25, 37, 50, 104 Grass, 1'22, 124, 145, 184, 189, 193, 196, anterior loop of, 24 208 intestinal, 25 reseeding with, 146 lateral oesophageal, 50 Grassland, 123, 144, 145, 159, 161, 194, valves, 25, 104 205, 206, 211 Heat extraction, 121-2 earthworms, as pests of, 223 Heath, Cal/una, 145 Gravelly loam, populations in, 163 Heathland, 144, 233 Gravelly sand, populations in, 162 Heavy metals, effects on worms, 220--1 Great Britain, 229, 248 pattern of uptake, 220 Green ash, 198 Hebrides, 163 Greenhouses, 95 Hedgehogs, 147 Greenland, 59 Heptachlor, 214, 218, 223 Gregarines, 148 Heptachlor epoxide, 218 Growing zone, 85 Herbage, 127 GENERAL INDEX 323

Herbicides, effects on worms, 205, 211, 100,104,113,114,152,168,184,187, 212 215, 220, 224 Herefordshire, 4 I dorsal wall of, 22 Hermaphrodite, I, 8, 36, 40, 76 Intra-epidermal capillaries of L. terres/rtS, Hertfordshire, 124 26 Heteroxanthine, 103 Introduced species, 63 Himalayas, East, 48 Invertebrates as predators, 147 Holland, 95, 124 Iron, 220 Holonephridia, 30, 3 I, 48, 57, 58 Irridescence, 14, 247 Horizontal distribution, 127-8, 242 Irradiation, 2 I 9 migration, 128 Irrigation, artesian, 206 Hormones, 106 Irritants, 109 Horse droppings, as food, 72, 231 Isobenzan, 2 I 4 5HT,106 Isotopes, 2 I 9 Human excreta, 185 Israel, 58, 59, 60 faeces, 228 Italy, 58, 59, 64, 66 urine, 228, 229 Ivory Coast, 144 Humidity, 19, 242 Humification, 172-4 Japan, 48, 58, 59, 64, 66, 73, 137, 138, 222, Humus, 159, 174, 193 229 Hungary, 58 Jaundice, 229 Hydrogen sulphide, 95 Java, 48 Hydrostatic pressure, 110 Jordan, 59 Hypotonic urine, 110 Jurassic period, 41

Kale, 166 Iceland, 59, 60, 147 Kentucky, 41 Immature worms, proportions of, 125 Impotency, 229 Kenya, 54 Kidneys, 101 Incorporation of fallen leaves, 168 'Kilner' jars, 23 I Inclusions, 101 Kommetjies, 142 inorganic, 19 Incubation period, 73, 74, 75 India, xv, 48, 49, 50, 5', 52, 53, 57, 58, 59, Lactic acid, 97, 98 60,65,66,73, 136, '4', '42, 144, 181, Lake District, 163 222 Lamellae, 34 Indiana, 124 Larch, 166 Indonesia, 57 Lashing movement, 88 Infectivity of cysts, effects of worms on, Lateral giant fibres, 34, 105 228 nerve cells, 34 Iriprganic minerals, '42 neural vessel, 18, 23, 24, 104 salts, 127 oesophageal hearts, 25, 50 Insecticides, effects of, 211, 242 oesophageal sub-neural vessel, 25, 26 residues, 2 I 5 oesophageal vessels, 23, 24, 25, 104 to control earthworms, 223 sub-neural commissures, 25 Inter\nediate hosts, earthworms as, 225 vessel, 23, 24, 25, 104 Internal salt concentration, I I I Latex, 139 Intersegmental grooves, I, 8, '3, 18, 248, Lawns, 223 25 0 Lead, 220, 22 I §Wtum,103 Lead arsenate, 2 I I zones, 13, 25 I Leaf, apple, 170 Intestinal epithelium, 99 ash, 237 microflora, 174, 186 bean, 237 wall, 33 beech, 170, 237 worms, 99 beet, 237 Intestine, 18, 20, 21, 22, 24, 48, 59, 98, 99, birch, 170, 237 324 BIOLOGY OF EARTHWORMS

Leaf-conI. layer, 122 burial, 236 pine, 170 cabbage, 237 rate of burial, 169 disks, 109, 165, 169, 170, 236 species, 98 elm, 170, 173, 237 turnover, 172-4 fall, 161, 168, 173, 235 Little Barrier Island, 53 kale, 237 Liver, 101 lime, 170, 237 Loam, 129, 162, 163, 231, 237 litter, 89, 90, 98, 108, 127, 165, 169, 170, Locomotion, 110, /[2-14 173 Lonbok,48 mineral content of, 165 Longitudinal canals, 30 oak, 170, 237 muscle, 4, 15, 17, 18, 22, 26, 34, 36 palatability of, 236-7 ridges, 13 protein content of, 166 vessel, 22 ribs, 170 Lucerne, 193 sugar content of, 166 Lumen, 27, 29, 30, 101 veins, 170 Luminescence, 95 weathering, 169 Luminous slime, 95 Leaf burial, effects of temperature on, 156 Lung worms, 225, 226 Leaf disks, rate of breakdown of, 170 Lymph glands, 24, 101 Leeches, 40, 41, 147 Lymphocytes, 19 Legume, 206, 207 Lettuce, 166, 222 Madagascar, 50, 5 I, 52, 53, 57 Leys, 176, 194, 206 Magnesium, 180, 201 Liberia, 50, 54 chloride, 113 Life cycles, 68-75, 239 sulphate, 119, 232 history studies, 239 Malagasy, 56 span, 72, 73 Malathion, 214 Light intensity, 93 Malay Archipelago, 48, 52, 56, 57 means of detecting, IS Peninsula, 57, 66 photonegative response, 107 Malaya, 49 photopositive response, 107 Malaysia, 66 reactions, 106-7, 178 Male cell, 37, 75 receptors, 106, 107 duct, 38 sensory. cells, 106 pore, 5, 8, 39, 44, 47, 49, 53, 55, 57, 58, stimuli, 107 59, 60, 75, 76, 78, 245, 249, 255 ultra-violet, 107 Mammals, 147, 215, 220, 227 Light loam, populations in, 162, 163 Manchuria, 66 Lignin, 177 Mangolds, 161 Lime, 170, 177, 197, 201, 208, 210, 211, Manure, 70, 93, 188, 200, 205, 239 234,237 Maoris, 229 Lipase, 99 Marginal cells, 27, 28 Lithium, II6 Maryland, 136 Litter, 89, 90, 98, 108, 122, 127, 165, 168, Mass migration, 94, 95 169, 170, 17 1, 172, 173, 174, 176, Mating 75--83, 81, 106 177, 178, 180, 189 Maze, 90, 92, 93, 95, 166, 206, 229, 243 breakdown of, 168-72, 178, 210 Mechanical, damage, effects on population, CN ratio of, 177-8 204 consumption, 172-4 stimulation, 122 decomposition, 178, 189 Mechanized soil washing, 119 feeders, 146 Medium loam, population in, 162, 163 forests, 165 Meganephridia, 30, 54 fragmentation of, 168-72 Megascolecid group, xi hazel, 173 Membrane, basal, 17, 33 humification, 172-4 basement, 16 incorporation of, 168 mesenteric, 18, 22, 23 GENERAL INDEX 325

Membrane--cont. Mountain ranges, 61 peri trophic, 100 Mouth, 2, 12, 20, 21, 31, 110 Menazon, 214 Mowrah meal, 120 Mercuric chloride, 120 Mozambique, 54, 57 Meronephridia, 30, 3 I Mucoproteins, 175 Mesh bag, 169, 170, 236, 237 Mucous cells, 14, 38, 39 Metabolism, effects of temperature on, 155 glands, 26, 96 Metallic residues in soil, 220 membrane, 114 Metham sodium, 211 tubes, 78 Methyl bromide, 2 I I Mucus, 14, 76, 94, 99, 100, 108, 138 Mexico, 51, 52, 53, 55, 57, 65 Mud,75 Michigan, 136 Mulching, 205 Microbial activity, 84, 174, 185, 187 Mull soil, 123, 145, 163, 192 cell, 186 species in, 123, 145, 163 protein, 175 Multiplication rate, 128 Microflora, 159, 168, 174, 182, 186, 187 Muscles, arciform, 76 Micronephridia, 30, 54 circular, 14, 15, 17, 18, 22, 26, 31, 32, Micro-organisms, 142, 168, 174, 182-9, 34, 36, 38, 112, 114 184, 185, 186, 187, 193 dorsal, 34 and earthworms, 182-9 fibres, 18 dispersal of, 188 innervation, 105 numbers in casts, 182, 184 longitudinal, 4, IS, 17, 18, 22, 26, 34, numbers in gut, 183 36 Middle tube, 27, 29, 102, 103 setal, 4 Mid-dorsal line, 18, 3 I sphincter, 10, 18, 19, 30 -ventral line, 32 subcuticular, 13 Migrations, 128, 131 ventral, 17 Millet, 196 Muscoid flies, 149 Mineral elements, 160, 181 Muscular configurations, 113 inorganic, 142 contractions, I 13 nutrients, 180-1 rhythm, II3 particles, 22, 190 tube, 27, 29 soil, 181 Mycobacteria, 187 Minerals, effect of, 160 Mineralization of nitrogen, 174-7, 209 Naphthalene, 169 Mississippi, 64 Narrow tube, 27, 28, 29, 102, 103 Mites, 174 Nematodes, 19,98, 148, 21 I, 215 Mixed woodland, 124 Nephridia, I, 10,21,26,27,28,29,3°,31, Moisture, 83, 127, 131, 132, 152 -4, 159, 48, 54, 55, 57, 58, 100, 101, 103, IIO 163, 195, 242, 243 enteronephric, 30 content of earthworms, 106, I I I, 112, exonephric, 29, 30 152 funnel, 27, 28 content of soil, 68, 109, IIO, II8, 121, holonephridia, 30, 3 I, 48, 57, 58 127, 131, 132, 133, 141, 152, 153, 159 loops, 30 equivalent, 201 macroic, 55 gradient, 154, 243 meganephridia, 30, 54 reaction to, 242, 243 meronephridia, 30, 3 I Moles, 147, 222, 223 micronephridia, 30 Molybdenum, 181 septal, 21 Moorland soil, 123 tubes, 27, 28 Morphology, xvi, 1-39 tufted, 30, 154 Mor soil, 123, 124, 145, 163, 192 vessel, 26 species in, 123, 163 Nephridiopore, 9, 10, 26, 27, 30, 44, 46, Mortality, 86 57, 58, 100, 103, 110 Motor fibres, 105 Nephrostome, 27, 28, 29, 101, 102, 103 nerves, 105 lower lip, 28 326 BIOLOGY OF EARTHWORMS

N ephrostome-cont. Novocaine, 106 peritoneum, 28 Number of samples, 122 upper lip, 28 Nutrients, 100, 180-1, 195 prostomial, 31, 32 Nutrition, 70 Nerve cells, 34 Nylon net, 166, 169 contralateral, 105 cord, 18,31, 32, 34, 35, 36, 37, 86, 104, 107, "3, "4, 116, 242 Oak, 124, 144, 166, 168, 169, 177, 184, 198 depressant, I 16 leaf litter, 169 fibres, 16, 105, 108 woodland, 124, 144 impulses, 108 Oats, 169, 197 intra-epidermal, 105 Obligatory diapause, 69, 83, 84, 242 lateral, 34 Observation cage, 237, 238 motor, 105 Oesophageal crop, 58 plexus, 15, 32, 33, 34, 35 hearts, 25, 50 prostomium, 32, 33 pouches, 20 prostomial, 31, 32 region, 18 ring, 32, 33, 34, 35 sacs, 2I segmental, 31, 32 wall,20 septal, 32, 33 Oesophagus, 20, 22, 24, 37, 45, 59, 98, 99, tissue, 14 104, 113 Nervous system, 31-5, 31, 32, 104, /05-9, Ohio, 149, 152 116 Oocytes, 36, 76 Netherlands, 128, 211 Oogenesis, 76 Nettle, 166 Oogonia,76 Neural lamella, 34 Oothecae, 68, 74 lateral vessel, 18, 23, 24, 104 Optic organelle, 16, 35 Neurofibrils, 16, 35 Optimum temperature, 116, 133, 157, 239 Neuroglia, 34 Orchards, 109, 122, 123, 124, 168, 171, Neurones, association, 105 173, 211, 214, 216 Neuropile, 34, 35 Ordovician period, 4 I N eurosecretions, 106 Organ, bush-like, 30 New Caledonia, 51, 52, 67 excretory, 10, 100, 101 New Guinea, 229 genital, 36, 40 New South Wales, 49, 59, 60, 172 gill-like, 26 New Zealand, xv, 49,50,51,52,53,59,67, photoreceptor, 35 84, 138, 139, 181, 195, 197, 201, 229 reproductive, 26 Newly-emerged worms, 85 respiratory, 26, 98 Nigeria, 50, 54, 55, 57, 75, 123, 136, 144, sense, 14, 35, 105, 108 173, 181, 195, 206 tactile, 109 Nile Valley, 144 Organic matter, 93, 98, 99, 127, 142, lSI)- Nitrate, 175, 176, 180, 189, 201 161, 164, 168-81, 183, 190, 192, 194, of soda, 160, 175, 209, 211 199, 201, 203, 205, 210, 21 1,235, 237 Nitro-chalk, 208, 209 breakdown, 168-81, 190, 205 Nitrogen, 102, 174-7, 180, 181, 189, 201, cycles, 168-81 202, 209 decomposition, 168, 176, 183 available, 180 effect of, 2 I2 content of L. terrestYlS, 177 incorporation of, 235 excretion, 27, 100, 176, 179 surface mat of, 159, 168, 172, 2 II from earthworm corpses, 175 Organo-chlorines, 214, 215-18, 217 humification, 174-7 Organo-phosphates, 218 mineralization, 174-7, 209 Osmotic pressure, I 10 non-available, 176 of blood, 110 production, 180 of urine, 101, 102 turnover, 176 regulation, I I I Noradrenalin, 106 Ova, I, 68, 76, 79, 81 GENERAL INDEX 327

Ovarian funnels, 36 Petroleum ether, 187 Ovaries, :n, 36, 37, 48, 49, 54, 58, 59, 76, pH, 99, 108, 109, 120, 127, 14g---52, 150, 81, 117 159, ISo, 181, 195, 201, 210, 234, 243 Oviduct, 21, 36, 37, 54, 76, 81 effects of, 150, 151,234 Ovisacs, 36, 48, 76, 81 reaction to, experiments, 243 Oxalic acid, 108 Phalaris leaves, 164 Oxidation-reduction potential, 159 roots, 164 Oxygen, 26, 96, 97, 98, 116, 159, 176, 178, Pharyngeal ganglion, 33 179, 186, 187, 189 gland, 20, 99 consumption, 98, 176, 179, 186, 187, 189 intestinal plexus, 35 debt, 97, 178 nerve plexus, 35 partial pressure of, 98 Pharynx, 20, 21, 24, 31, 98 tension, 159, 178 Pheromone, 140 Philippines, 48, 66, 222 Pacific Islands, 49 Phorate, 214, 223 Pakistan, 48, 59, 60 Phosphorescence, 95 Palearctic zone, 47 Phosphoric acid, 108 Palatability, 161 Phosphorus, ISo, 181, 201, 210 experiments, 236-7 Photoreceptor cells, 15, 16, 107 of leaf litter, 108, 109, 161, 166 Photosensitivity, 107 Panama, 50, 56 drug depressants, 107 Papillae, glandular, 8 Physiological experiments, 242 Paraguay, 52, 56 Physiology, xvii, 96-II7 Paraquat, 212 Pig, kidney of, 227 Parasites, 19, 147-9, 223, 225, 226, 227, litter, 124 228 Pigment migration, 106 eggs of, 228 Pigmentation, 13-14, 84, 85 nematode, 19 Pigs, 225 protozoan, 19, 148 Piles, 229 transmission of, 226, 227 Pine litter, 170 Parathion, 116, 2 I 4 needles, 165 Parthenogenesis, 63, 64, 76, 79, 8 I, 242 woodland, 124 facultative, 79 Plant breakdown, xvi Passive agents, earthworms as, 228 cover, 159 Pasture, 84, 123, 124, 129, 133, 136, 138, growth, 181, 195, 206, 228 144, 151, 156, 159, 161, 165, 169, 172, material, 164, 207, 231 181, 188, 190, 191, 193, 194, 197,201, pathogens, earthworms as vectors of, 188 206,207,208,217, 231, 232, 233, 234 residues, 193 productivity, effect of worms on, 198 Ploughing, 145, 151, 177, 203, 204 Peas, 196 effects on worms, 2 I2 Peat, 72, 237 Plugs, burrow, 89 Penes, 78, 8 I Podsol, 123 Peregrine species, 49, 61, 64, 66, 246 Polyhydric phenol, 166 Perichaetine setae,S, 245 Polynesia, 52 Peri-enteric plexus, 18, 25, 104 Polyphenols, 108, 166, 169 Peristalsis, II 2-14, II3 water soluble, 166, 169 Peristomium, 2, 60 Polysaccharide gums, 188, 193 Perithecia, 188 Population, Peritoneal cells, 19, 101 adolescent, 87 Peritoneum, 14, 15, 17, 25, 28, 34, 76 biomass, 206 Peritrophic membrane, 100 density of, 162 Persia, 59 distribution of, 127-31 Peru, 51 Population, estimation of, 118-23, 233 Pesticides, effects of, 2II-15 factors limiting, 177 uptake in earthworms, 215-18 geographical, 6I -7, 203 Pests, earthworms as, 222-30 horizontal distribution, 127, 128, 242 328 BIOLOGY OF EARTHWORMS

Population-con/. Proteolytic enzymes, 99 numbers, 162 Protozoa, 98, 99, 148, 188, 226 pyramid, 74 Protozoan parasites, 19, 148 reduction of, 147 Pseudotsuga mor, 124 seasonal activity, 131-8, 134 seasonal changes in, 87, IJI-8, 132, 133, QlIadrat, II8, 121, 123, 234, 235, 242 136, 137 Quaternary ice caps, 64 size of, 12J-7 Queensland, 49, 5 I, 52 structure, 125-7 Quiescence,8J-4, 130, 13 1, 136, 153 vertical distribution, 128-31, 129, 138, Quinine, 17 242 Populations, 118, 127, 128, 136, 138, Radiation, 95 147, 151, 152, 156, 160, 161, 162, 164, effect on A. chlorotlCa, 219 177, 178, 179, 189, 195, 203, 205, 206, gamma, 219 207, 21 4 Radioactive fallout, 2 I 8 Pores, cuticular, 14 pollution of soil by, 2 I 8 dorsal, 8, 19, 20, 26, 55, 57, 58, 60, 88, Radiocaesium, 173, 219 IIO, 151,248,249,250,251,252,253, Radio-iron, 219 256 Radioisotopes, effect on worms, 2I 8-1 9 female, 8, I I, 48, 49, 55, 57, 58, 60 uptake into worms, 218, 219 genital,s, I I Radishes, 228 male, 39, 44, 53, 57, 58, 59, 60 Ragi,228 prostatic, 8, 42, 44 Rain, reactions to, 95 spermathecal, 8, 48, 53, 60, 245, 247 Rainfall, 235 Porphyrins, 13 Raw humus, 159 Porosity, 194-5 Reception cells, 114 Post-clitellar region, 6 zone, 99 Post-septal canal, 27 Receptors, 17, 35 Pot experiments, 197, 213 Rectum, 195 Potassium, 180, 181, 210 Reduction division, 76 cyanide, II 6 Regeneration, 80, 85, Il4-16, 243 permanganate Regenerative phenomena, 80, 106, lIS, Potato root eelworm, 224 II6 Pouches, 18 Rennin, 99 Predation, 70 Replacing cells, 14 Predator, 20, 143, 147-9 Reproduction, 75-8J, 106 Pregnancy, testing, 229 asexual, 79, 80, 8 I Preseptal canal, 27 effects of temperature on, 154 Preservation, 2JJ Reproductive cells, 75 Proboscis, 2 cycle, 106 Prolobous prostomium, 2, 4 potential, 127 Propham, 212 structures, 21 Proprioceptor cells, 35 system, 36-9, 36, 37, 60 Prostates, 8, 21, 39, 44, 45, 49, 53 Reservoir, 27 Prostatic fluid, 78 Reservoir host, earthworms as, 225 gland, 46, 48 Resorption, 101, 102 pore, 8, 42, 44 Response, learned, 94 Prostomial nerve, 31, 32 Respiration, I, 26, 96-8, I 16, 178, 180 Prostomium, 2, 4, 12, 15, 17, 31, 32, 33, anaerobic, 98 57, 60, 95, 106, 108, 110, 112, 245, cutaneous, 20 248, 249 depression of, 98 enervation, 3 I, 35 effects of temperature on, 154, 155 Protease, 99, 142 rates of, 97, 98 Protective slime, 95 Respiratory exchange, 187 Protein, 101, 102, 103, 166, 175 system, 26-7 Protein content of earthworms, 229 Resting phase, 84 GENERAL INDEX 329

Retinella, 35 nerves, 31, 32 Rheumatism, 229 Segmentation, 1-4, 239 Rhizopods, 184 internal, I Rhodesia, 54, 57 Segments, I, 8, 27, 105, 109, 151 Rice, 222 Selenium, 220 Ridges, 12 Semi-aquatic species, 9, II, I II, 154 Rigor, 110 Seminal fluid, 78, 79 Rio de Janeiro, 56 grooves, 8, 12, 76, 247 Rodents, 226 recepticles, 8 I Rolling, effects of, 203 vessels, 36, 37 Roots, 161, 164, 165, 168, 193, 198, 200, Seminal vesicles, 21, 37, 57, 58, 60, 79, 81, 205, 222 148,230 Rotary cultivation, 203 anterior, 37 Rotations, 206 mid,37 Rothamsted, 125, 126, 129, 133, 145, 151, posterior, 37 160, 161, 172 Senescence, 85 Rotifers, 98 Sense organs, 14, 15, 35, 105, 108 Rowan, 177 epidermal, 16 Russia, 58, 59, 60, 163 epithelial, 35 Rye, 184, 197, 206, 201) photoreceptor, 35 Sensory cells, 14, 15, 16, 17, 34, 106 Septal nerves, 32, 33 hairs, 35 wall,27 input, 106 vessels, 25 light receptive role, 106 Septum, 22, 27, 29, 32, 36 Septa, 1, 18, 20, 25, 30, 104, 112 intersegmental, 102 innervation of, 34 Setae, I, 4, 5, 6, 7, 41, 48, 53, 55, 57, 58, maturity, 12, 72, 86, 87, 155, 188, 242 60, 78, I12, 114, 175, 245, 248, 249, organs, 40 250, 25 1, 253 Seychelles, 50 closely-paired, 6, 7 Sheep droppings, as food, 164, 240 copulatory, 58 Siberia, 59, 60, 64 distant-paired, 6, 7 Sicily, 58, 66 genital,s, 6, 60, 8 I Sieve, 119, 121, 232 lateral, 12 Silurian period, 4 I lumbricine, 5 Simazine, 212 normal, of L. lerreslns, 5 Simple key, 245-53 penial,81 Size of sample, 122 perichaetine, 5, 245 Slime, 95 physical stimulation of, 5 luminous, 95 ventral, 12 protective, 95 widely-paired, 6, 7 tube, 76 Setal arrangements,s, 7, 245 Slugs, 147 lumbricine, 7 Smallpox, 229 perichaetine, 7 Snakes, 147 Setal muscles, 4 Sodium chloride, I I 1 Setal form, 4, 5 Soil, 98, 99, 109, 145, 149, 150, 151, 152, Sewage sludge, 70 154, 156, 159, 163, 168, 170, 181, 184, Sexual activity, 75 185, 200, 216 development, 76 acidity, 108, 149, 150, 151, 208 Secretory zones, 99 adverse effects on, 228-9 Seedlings, 222 aeration, xvi, 127, 190, 194-5, 205, 206, Segment numbering, 233 228 Segmental bands, 13 air space, 194 contractions, I 12 amelioration, 199-203, 229 ganglia, 31 animals, 168 grooves, 17 climate, 68 330 BIOLOGY OF EARTHWORMS

Soil-cont. forest, 123, 193, 194, 201 climate, fluctuations of, 68 garden, 194 compaction, 192 gravelly loam, 163 consumption, 172-4 light, 162, 163 degeneration, 2 I I light loam, 162, 163 density, 202 light sandy, 163 drainage, 194-5, 205, 206, 228 medium loam, 162, 163 erosion, 192, 228, 229 mor, 123, 124, 145, 163, 192 fertility, xvi, 174, 181, 190-2, 199, 205 mull, 123, 145, 163, 192 flooded, 153 open, 162 formation, xvi, 200 orchard, 159, 171, 21 I fumigants, 21 I peaty, 145, 163 horizons, 192 ploughed, 180, 200 indicators, 195 sandy, 144 ingestion of, 190 shallow acid peat, 163 litter species, 98 tropical, 123 microbial activity in, 174 tropical, populations in, 123 microflora, 159 water-holding capacity, 190, 195 mineral fraction of, 191 water-logged, 109, I II, 156 minerals in, 18 I water-saturated, 153 moisture, 68, 109, IIO, II 8, 121, 127, woodland mull, 145, 163 13 1, 132, 13L 141, 15~ 153, 159 South Africa, 222, 229 movement through, 14 South America, 50, 5 I nutrient content, 195 South Shetland Islands, 67 organic matter, 161 Saccharose, 108 organisms, 174 Salamanders, 147 particles, 174, 190, 193 Salivary glands, 99 particles, breaking down of, 190 Salt, III pathogens, 21 I internal concentration, I I I pathogens, control of, 2 I I saturated solution, 242 pH, 200 Sample number, 122 profile, 168, 171, 192, 199 size, 122 reclamation, 199, 200 Sampling methods, 118-23, 233, 234, 242 stability, 211 chemical, 120-1, 131, 133 sterilized, 188 digging, 232 structure, xvi, 190-2, 194, 228 electrical, 122 surface, 188, 201, 21 I, 223, 234 formalin, u4, 232, 233, 234, 242 temperature, 68, 84, 97, 121, 127, 129, handsorting, 118-19, 233, 234, 242 13 1, 132, 133, 138, 155, 157, 179 heat extraction, 121-2 texture, 127 soil cores, 118, 122 throughput, rate of, 123 vibration, 122 turnover, 172-4, 191-2, 205 vibrating rod, 122 Soil type, 162-3, 191-2, 205 washing, I 18, 122 agricultural, 152, 228 wet sieving, u4 air filled, 153 Sampling tool, 118 alluvial, 162, 163 Sand, 190 brown earth, 181 Sardinia, 52, 58, 66 brown podzol, 196 Savannah, 66, 125, 195 clay, 162, 163 Scandinavia, 255 clay content, 161 Scotland, 145, 162 clay, effect of, 161 Seasonal abundance, u6, 128, 139 coniferous forest, 146 activity, 131-8 coniferous mor, u4, 145 changes, 126, 139 dark, 163 Secondary sex characters, 84, 106 Egyptian, 161 Secretory cells, 106 fallow, 124 Soybeans, 196, 207 GENERAL INDEX 331

Species, 42---{)0 Submergence, 154 associations, [45-7 Subneural vessels, 18, 23, 25, 26, 104 Specific gravity, 232 Suboesophageal ganglion, 31, 106 Sperm, 81 Subpharyngeal connectives, 33 cells, 39 ganglion, 32 funnels, 36, 37, 38, 48, 75 Substrate, 109 sacs, 38 Subsoil, 186, 190 transference, 42, 78 Subtillers, 205 Spermathecae,8, 13,21,36,37,39,42,48, Sucrose, 17, 108 54, 59, 60, 78, 81, 148 Suction pump, pharynx as, 20 Spermathecal ducts, 39 Sudan, 57 openings, 76, 78, 79 Sulphate of ammonia, 172, 21 I pores, 8, 48, 53, 60, 245, 247 effects of, 210 Spermatogenesis, 75, 230 Sumatra, 48, 66 Spermatogonia, 75 Sumithion, 214 Spermatophores, 8 I Sunda Islands, 48 Spermatozoa, 75, 78, 79 Superphosphate, 208, 210 Sphincter muscles, 10, 18, 19, 30 Supra-intestinal, glands, 57 Spores, 184, 186, 188 ventral commissures, 25, 26 fungal, 185, 224 vessel, 23, 24, 25, 26 Sports fields, 213, 223 Surface casting species, 131 earthworm control on, 213 debris, 216, 232 Spraying, 168 dwelling species, 121, 150, 204, 232 Springtails, 174 Survival, 152 Spruce, 166 in sugar solutions, 242 Squirter earthworm, 19 in test chambers, 242 Staphylinid beetles, 147 in water, 152, 242 Starlings, 147 Sweden, 131, 146, 161 Stems, 168 Swine, 226 Stewart Island, 51 lungs of, 226 Stimuli, 89, 90, 106 stomach of, 226 adverse, 105 Switzerland, 64, 144 avoidance of, 92 Symbiotic bacteria, 99, 182 chemical, 89, 90, 105, 108 Sympathetic system, 33 electrical, 89, 109, 199, 243, 244 Synapses, 105 electrical response to, 89, [09 Syria, 58, 59, 60 intersegmental, 112 Systematic affinities, 40-2 light, 76, 93, 107 light response to, 107 Tactile organs, 109 photic, 107 receptors, 109 tactile, 15, 88, 93 Tannins, 166 touch, 76, 109 I 12 Taste, 17 vibrations, 89, 95 Tanylobous prostomium, 2, 4, 249 Stomach, birds, 226 Tanzania, 54, 57 swine, 226 Tapeworms, 225 Stones in the bladder, 229 Tartaric acid, 108 Straw, 70, 174, 197, 23 1, 239, 240 Tasmania, 51 Stress, 20 Taxonomy, 40---{)7 Stretch receptors, 35 TeA, 212 Stubble-mulch farming, 205 Teleospores, 224 Stubble stripping, 204 Telodrin, 214 Sub-aquatic species, 43 Temperate forest, 173 Sub-Arctic Islands, 51 zones, 65, 97, 168 Subcuticular muscle, 13 Temperature, 68, 61), 70, 72, 83, 135, 141, Sub-epidermal nerve plexus, 33, 34, 105 179, [54-8, 158, 23 1, 235, 239 Sub-epithelial nerve plexus, 34 acclimation, 157 332 BIOLOGY OF EARTHWORMS

Temperature--cont. Tubificity, 41 conditioning, 157, 158 Tumours, 230 effect of, 244 Tunis, 58 effect on excretion, 103 Turf, earthworms as pests of, 223 effect on respiration, 97 Turnover of soil and litter, [72-4, [9[-2, gradient, 157 205 lethal, 156-}, 158 Typhlosolar vessel, 25 optimum, 116, 133, 157, 239 Typhlosole, 22, 25, 57, 104 preferenda, 156, 157, 158 seasonal changes in, 158 Uganda, 57, 123 Tension receptors, 35 Ubiquitous species, 150 Termites, 147 Ultra-violet light, 95, 98, 107 mounds of, 147 effects of, 95, 98 Testis, 37 Urea, 100, 101, 102, 103, 175 Testes, 21, 36, 48, 49, 59, 75, 79, 81, 148 Uric acid, 100, 103, 175 funnels, 59 cells, 101 holoandric, 36 Urine, 100, 101, 102, I II, 175, 176, 198, meroandric, 36 228, 229 Thermal death point, 157, 158 osmotic pressure of, 101, 102 Thigmotactic reactions, [09 U.S.A., 57, 64, 65, 66, 124, 127, 13 1, 133, Thiophanate-methyl, 214 136, 138, 147, 149, 152, 156, 180, 195, Thrushes, 147 199, 204, 206, 21 6, 229 Timothy grass, 206 U.S.S.R., 124, 200, 206 Tissue, connective, 17, 34, 38 nuclei of, 38 Vacuoles, 100 production, 177 Vagina, 228 'T' maze, 90, 92, 93, 94 Variance of species, 60 Toads, 147 of subspecies, 60 Tobacco, 222 Valerionic acid, 98 Togoland, 55, 57 Vas deferens, 21, 36, 37, 38, 39, 48, 53, 75, Tooth powder, 229 81 Topsoil feeders, 146 racemose, 39 Touch, 88 termination of, 53 Trachea, 227 tubular, 39 of bird and chickens, 226, 227 Vas efferens, 37, 38, 75 Transplantation, [[6-[7 Vascular system, I, 22---{) Transverse commissure, 107 Vesiculae seminalis, 3 I septa, 17 Vector diagram, 46 Triassic era, 41 Vectors of animal viruses, Triazine herbicides, 212 earthworms as, 225 Trichlorphon, 214 Vegetable crops, 222, 228 Tropical areas, 97 Vegetative cells, 184 forest, 173 Venezuela, 53, 56 lowlands, 64 Ventral muscle, 17 soil, 123 nerve cord, I, 23, 31, 32, 33, 34, 35, 37, Tropics, 136, 144 105, 107, 114 Tube, ciliated, 101, 103 vessel, 18, 23, 24, 25, 37, 103 middle, 27, 29, 102, 103 view, 12 mucous, 78 Vertebrates, 147 muscular, 27, 29 Vertical distribution, [28-JI, 129, 138, 242 narrow, 27, 28, 29, 102, 103 migration, 131 slime, 76 Ventral ganglion, 106 wide, 29, 102, 103 Vessel, afferent nephridial, 18 Tubercula pubertatis, II, 12, 13, 60, 81, afferent typhlosolar, 18 245,248,249,250,251,252,253,255 blood, 15, 18, 22, 26, 33, 38, Il6 Tubercules, 12, 252 capillary, 14 GENERAL INDEX 333

Vessel-cont. Weathering, 169, 191 commissural, 25 Weight, 86, 139 dorsal, 18, 21, 22, 23, 24> 25, 26, 37, 104 live, 123, 167, 173, 218 dorso-intestinal, 18, 25, 104 seasonal increases in, 86 dorso-subneural, 18, 23, 25, 104 wet, 218 efferent nephridial, 18 Westmorland, 124 lateral-neural, 18, 23, 24, 104 West Indies, 50, 51, 52, 53, 55, 56, 66, 67 lateral-oesophageal, 23, 24, 25, 104 Wheat, 160, 196, 206, 212 lateral-oesophageal subneural, 25, 26 Wide tube, 29, 102, 103 longitudinal, 22 Withdrawal reactions, 88 nephridial, 26 Woodland, 123, 124, 145, 156, 159, 161, subneural, 18, 23, 25, 26, 104 168, 169, 172, 173, 177, 179, 233 supra-intestinal, 23, 24, 25, 26 beech, 124 typhlosolar, 25 mixed, 124 ventral, 18, 23, 24, 25, 37', 103 oak, 124, 144 ventro-intestinal, 25, 104 species, 146 ventro-oesophageal, 104 Worm cairns, 89 ventro-parietal, 18, 25, 26, 104 Wound tissue, 114 Vibrations, 89, 95, 122 Wych Elm, 166 Victoria, 49, 5 I Viruses, 148 Yeasts, 184, 186 Voltage, 109 Yield increases, 195, 196, 197 Yolk, 40, 76 Wales, 124, 151, 220 Yucatan, 51 Water, 103, 152 conservation, 109, 110 Zambesia, 57 infiltration rates, 195 Zanzibar Island, 54 loss, IIO, 152 Zinc, 220, 22 I relationships, 109--I 2 Zinophos, 214 uptake, I10, 154 Zurich, 144 Water stable aggregates, 187, 190, 192, 193 Zygolobous prostomium, 2, 4, 57, 248