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BULLETIN DU MUSÉUM NATIONAL D'HISTOIRE NATURELLE

3 e série, n° 450, mars-avril 1977, Zoologie 313

Preliminary descriptions of Indian earthworms

(Megascolecidae : Oligochaeta) from the Palni Hills

by B. G. M. Jamieson *

Résumé. — Descriptions préliminaires de Vers de terre (Megascolecidae : Oligochaetes ) des col¬

lines de Palni ( Indes).

Huit espèces de Vers de terre des collines de Palni (Indes), dont l’une appartient à un nouveau

genre, sont décrites : Diporochaeta dorsochaeta n. sp., D. macrochaeta simpliciseta n. s.sp., Celeriella

bursata n. sp. et C. punctata, n. sp., C. quadripapillata Stephenson, 1924, Troyia gundarshola n. gen.

et n. sp., Lampito bouchei n. sp. et L. sylvicola Michaelsen, 1907. Les diagnoses des genres sont

données ainsi que des clés dichotomiques pour les espèces des genres Celeriella et Lampito. Toutes

les espèces orientales attribuées jusqu’à présent au genre Plutellus sont transférées dans le genre

Diporochaeta qui nécessite une nouvelle étude. Le genre Celeriella présente des rangées transver¬

sales de néphrostomes préseptaux au lieu de la paire médiane qui lui avait été attribuée précédem¬

ment.

Abstract. — Eight species of earthworms from the Palni Hills including a new genus, are

described : Diporochaeta dorsochaeta sp. nov., Diporochaeta macrochaeta simpliciseta suhsp. nov.,

Celeriella bursata and C. punctata spp. nov., C. quadripapillata Stephenson, 1924, Troyia gundar¬

shola gen. et sp. nov., Lampito bouchei sp. nov. and L. sylvicola Michaelsen, 1907. Their respective

généra are delined and keys to the species of Celeriella and Lampito are given. Ail Oriental species

formerly assigned to Plutellus are transferred to Diporochaeta but the latter genus requires further

analysis. Celeriella is shown to hâve transverse rows of preseptal nephrostomes, not the médian

pair previously attributed to the genus.

Introduction

Ecological studies conducted by Professor J. P. Troy in the Palni Hills of South India

hâve yielded nine species of earthworms of which one, Moniligaster troyi is described elsewhere

(Jamieson, 1976&) and eight are the subject of this paper. It lias heen concluded (Troy,

in litt.) that earthworms play a very important rôle in soil-forming processes of these

hilly tracts, in their fertility and even in the stability of their landscapes. Description

of these worms is therefore of value not only to taxonomists abut lso to ecologists and

pedologists working in the area.

The earthworms were collected al five stations from longitude 77°16'27" and 77°28'05" E

and latitude 10°08'21" and 10°12'49" N in the Palni Hills. Detailed descriptions of the

habitat will be presented by Professor Troy in a fortbcoming publication but abbreviated

descriptions are given here.

* Zoology Department , Universily of Queensland, St Lucia, Brisbane 4067, Australia.

450, 1

478

B. G. M. JAMIESON

COLLECTING LOCATIONS IN PaLNI IIlLLS

Station 1. — South India, border between Kérala State and Tamilnadu, 77° 16'27" E 10° 08'21" N,

altitude 2 490 ni, Vandaravu Range, topography hilly, summit of a local dôme. Température :

mean annual 13°C, amplitude (annual) 4-5°C, daily up to 40°C within the grass eover ; frost

frequent ; rainfall annual average 1 400 mm ; fog frequent ; dry season 2-3 months (December-

February). Végétation Pollinia phaeotrix and Arundinella vaginata -f- hygrophytes, low

savanna (montane grassland), probably climax ; trees and shrubs absent from grass cover ;

fire regular each year (January, February). Rock types intermediate to acid charnockites

traversed by quartz and pegmatite veins, and enderbite. Soil montane humic ferrallite

(with highly organic upper horizons, up to 25 % organic matter in top soil, well associated),

typic gibbsihumox humic ferralsol ; pfl (upper horizon) 4.9 ; water 1/2.5.

Station 2. — Talmilnadu, 77° 25'46" E 10° 11'08" N, altitude 2 350 m, top of Southern greatclifî,

along road Kodaikanal-Munnar, 15 km WSW of Kodaikanal, topography hill slope (30 %).

Température : mean annual 16°C, amplitude annual 3-4°C, daily up to 50°C within the grass

cover ; frost frequent ; rainfall annual average 1 400 mm ; fog moderately frequent ; dry

season 3-4 months (December-April). Végétation Chrysopogon zeylanicus and Arundinella

çaginata savanna, with various small trees and shrubs (typical montane savanna) ; fire regular,

each year during dry season. Rock type intermediate to acid charnockite. Soil montane

intergrade between humic ferralite and ferralitic with gibbsite soils ; highly organic (up to

30 % organic matter) top soil, well decomposed and associated, typic humic gibbsihumox,

humic ferralsol ; pH (upper horizon) 5.3 ; water 1/2.5.

Station 3. —- Tamilnadu, 77° 26'37" E 10° 12'49" N, altitude 2 270 m, central Palni-Gundar Rasin,

3 km W of Kodaikanal Observatory, topography moderate hill slope (18 %). Température :

mean annual 15°C, amplitude annual 3°C, daily < 10°C under tree cover ; rainfall annual

average 1 300 mm ; dry season 3-4 months (December-April) ; frost nil under tree cover.

Végétation : old blue gum plantation (1920), Eucalyptus globulus, pure dense. Rock type

intermediate charnockite (highly metamorphic hornblende-hypersthene gneiss). Soil montane

humic ferrallitic soil with gibbsite with moder-mor top horizon, humic gibbsihumox, ferral¬

sol ; pH (top horizon) 4.4 ; water 1/2.5

Station 4. — Tamilnadu, 77° 28'05" E 10° 12'21" N, altitude 2 285 m, Kodaikanal golf links near

pillar rocks, 5 km S of Kodaikanal ; topography gentle slope. Température : annual mean

16°C, amplitude annual 3-4°C, daily up to 15°C at soil level ; rainfall annual average 1 300/

1 500 mm ; dry season 2-3 months ; frost possible under trees during cool season. Végétation

old pine plantation (exotic), Pinus radiata (1920) with understorey of exotic Acacias, A. deal-

bata, A. mearnsii, A. melanoxylon ; cover 0.6-0.7. Rock : jointed (orthogonal joints) inter¬

mediate charnockite. Soil montane ferrallitic with gibbsite soil with moder-mor top horizon,

humic gibbsihumox, ferralsol ; PH (upper horizon) 4.1 ; water 1/2.5.

Station 5. — Tamilnadu, “ Gundar Shola ”, Gundar Basin, Central Palni, 77° 26'13" E 10° 12'46" N,

altitude 2 100 m ; topography steep slope, northern aspect. Température : annual mean

15°C ; amplitude annual 2-3°C ; daily < 10°C under forest ; rainfall annual average 1 300-

1 400 m ; dry season 3-4 months ; frost nil under tree cover. Végétation natural forest,

montane evergreen low forest dominated by Lauraceae, Elaeocarpaceae, Myrtaceae, Icaci-

naceae, etc. ; very little disturbed. Rock type intermediate to acid charnockite. Soil forest

montane ferrallitic with gibbsite soil, with well associated organic matter to a depth of 120 cm,

humic gibbsihumox, ferralsol ; pH (upper horizon) 6.3 ; water 1/2.5.

MEGASCOLECIDAE FROM THE PALNI HILLS

479

Distribution of species

(+ = présent)

Moniligaster troyi .Tamieson, 197 6b

Diporochaeta dorsochaeta sp. nov.

Diporochaeta macrochaeta slmpliciseta subsp.

Celeriella bursata sp. nov.

Celeriella punclata sp. nov.

Celeriella quadripapillata Stephenson, 1924

Troyia gundarshola gen. et sp. nov.

Lampito bovchei sp. nov.

Lampito sylvicola Michaelsen, 1907.

Stations

nov.

Tribe PERIONYCHINI Jamieson, 1971a

Genus DIPOROCHAETA Beddard, 1890

Setae 8 or more per segment. A pair of combined pores of vasa deferentia and tubular

or tubuloracemose (rarely racemose ?) prostates on XVIII. Last hearts in XII or XIII,

typically those in XII posteriad latero-oesophageal ; subneural vessel absent. Gizzard in

V or VI (well developed to vestigial) ; typhlosole absent or exceptionally présent (rudi-

mentary to well developed). Extramural caleiferous glands usually absent, rarely with

paired glands in X and XI (type-species) or XI and XII or unpaired annular glands in XV.

Nephridia stomate, exonephric holonephridia with or without bladders ; their pores in straight,

or sinuous lines but never with regular alternation ; the anterior nephridia sometimes

tufted ; rarely ( dorsochaeta ) with two at least caudally stomate nephridia on each side per

segment. Spermathecae 1-5, pretesticular, diverticulate, paired or rarely ( palniensis)

médian unpaired ; diverticulum usually single, uniloculate, rarely paired, exceptionally

composite, never sessile multiloculate.

Distribution : New Zealand (type-species) and neighbouring islands. Eastern Subregion of

Australia. India. Ceylon. Burma.

Type species : Perichaeta intermedia Beddard, 1889.

Remarks

The above brief diagnosis of Diporochaeta is modified from Jamieson (1976a), where

a detailed description and a discussion of the genus are given, to allow inclusion of oriental

species hitherto assigned to the purelv Australian genus Plutellus. The sole distinction

between the classical Plutellus and Diporochaeta, the perichaetine condition of the latter

and the lumbricine condition of the former, has been shown in the work cited toi ack signi-

ficance. American “ plutelli ” are referable to Argilophilus Eisen which is not clearly

450, 2

480

B. G. M. JAMIESON

separable from Diporochaeta but appears to difïer notably in the absence of extramural

spermathecal diverticula. Incorporation of oriental species, including the two aberrant

species dcscribed below [D. macrochaela and D. dorsochaeta ) lias necessitated the emend-

ments to the generic diagnosis for the following features : the typhlosole may be well deve-

loped ; nephridia are rarely replicated [D. dorsochaeta) ; and spermathecae may be unpaired.

Restriction of Plutellus to species which, like the type-speccis, hâve thethree apomorph

characters well developed extramural calciferous glands in X-XIII in combination witli

vesiculate nephridia and regular alternation of nephropores (Jamieson, 19716) lias been

formally recognized by Jamieson and Nash (1976). While such restriction appears wholly

justified, it must be noted that Diporochaeta (with its possible junior synonym Graliophilus)

is defined largely by the synplesiomorphy liolonephridia with no regular alternation of

nephropores (and in most species tubular prostates) and it possibly will be divisible into

further généra by application of the principle of Hennig that taxonomie groups should

be recognized on the basis of shared advanced characters (synapomorphies). Such an

analysis of Diporochaeta has in efïect been commenced by removal of D. pellucida Bourne

Nilgiri Hills) to the monotypic Priodochaeta by Gates (1940). P. pellucida resembles D. macro-

chaeta in having tufted nephridia anteriorly and liolonephridia caudally but differs in the

apomorphy extramural calciferous glands (in XIV-XVI). The two species may be more

closely related than they are to other members of the Diporochaeta-Priodochaeta complex.

A key to the previously known species of oriental Diporochaetas is given, under Plu¬

tellus, by Gates (1972).

Diporochaeta (?) dorsochaeta sp. nov.

(Fig. 2 B, C ; 5 E ; 6 E-G, K ; Table I)

Length 39,66 mm, width (midclitellum) 1.1, 1.4 mm, segments 110, 118 (H, PI). Pro-

stomium epilobous, almost tanylobous, sides slightly convergent posteriad ; hind end clubhed.

First dorsal pore 5/6. Setae commencing in II in 8 regular longitudinal rows throughout ;

setae d displaced far dorsally, closer together than they are to c lines in ail segments and

at the two ends of the body conspicuously enlarged (FI, PI) ; in XII aa : ab : bc : cd : dd

averaging 1.9 : 1.0 : 1.8 : 2.1 : 1.2 ; dd : u = 0.07-0.13 (mean 1.2, 8 specimens). Nephro¬

pores not externally recognizable. Clitellum annular 1/2 XIII-XVI, weakly (H) to well deve¬

loped 1/2 XIII-XVII (P2) ; at full development occluding intersegments and dorsal pores ;

setae retained. Male pores in b lines of XVIII relative to adjacent segments, on stron-

gly protubérant papillae which greatly expand the setal annulus but leave free anterior and

posterior bands of the segment ; each papilla précédée! by an indistinct oval génital marking

which reaches 17/18 (H, P2) or this marking not apparent (PI). Female pores a pair

anteromedian of setae a of XIV. Spermathecal pores 2 pairs of minute orifices, in 7/8 and

8/9, in b lines (H, PI).

Thickest septa 7/8-10/11, only moderately thickened (H). Dorsal blood vessel single

not certainly traceable on and in front of the gizzard (IT, PI) ; last hearts in XII ; those

in X-XII arising from the supra-oesophageal and apparently also from the dorsal vessel

(latero-oesophageal) ; supra-oesophageal small, in X-XII ; subneural vessel absent. Giz¬

zard in VI, large and strongly muscular, globose (H) to eylindrical (PI) extending into VIII

MEGASCOLECIDAE FROM THE PALNI HILLS

481

by backward deflection of septa. Oesophagus lacking extramural calciferous glands but

widened and with numerous high longitudinal radial lamellae in XIII and XIV (H) or

XIV and XV (PI) or XIII, XIV, XV (P2). Intestine coinmencing in XVII (H), 1/2 XVII

(PI), XVIII (P2) or XIX (H) if the portion in XVII and XVIII, which is more vascular

than in XIX, be considered oesophageal ; an internai constriction anteriorly in XVII is

apparently the oesophago-intestinal valve, however ; muscular thickening caeca and typhlo-

sole absent. Nephridia : in the anterior intestinal région with two large nephridia on eaeh

side, discharging by a long slender duct presetally in cd and b respectively ; the latéral

nephridium with a (preseptal ?) funnel ; no fonnel demonstrated for the médian nephri-

dium. Two nephridia on each side in the forehody also. Caudally with this condition

persisting but the latéral, stomate nephridium much enlarged ; médian preseptal funnel

demonstrated for most caudal segments and in some segments a further latéral funnel,

suggesting a simple Celeriella condition. Holandric ; iridescent funnels in X and XI ;

testis-sacs absent; large, much divided séminal vesicles in IX and XII. Prostates

one pair, depressed tubular in XVIII, the gland much contorted into a compact

rounded or almost quadrangular mass, depressed and irregular in cross section chiefly

owing to mutual distortion of adpressed coils ; ectal duct slender and tortuous, though

muscular ; vas deferens joining the entai end of the duct (H, PI). Penial setae présent :

short and stout, almost tusklike, tapering to blunt points and with a few small apically

directed teeth sparsely scattered over the apical sixth ; a mature seta 237 [xm long and at

inidlength, 11 um wide. Ovaries, with several terminal strings of large oocytes, and funnels

in XIII ; small loculi in XIV may be ovisacs. Spermathecae two pairs in VIII and IX,

each retort-shaped with ovoid ampulla and slender duct (H, P'l), the ampulla recurved

on the duct in the holotype ; a short clavate diverticulum (inseminated in Stn 2 material)

pendant laterally near the entai end of the duct.

Material examined : Stn 5 — holotype (Paris Muséum AH 320) ; paratypes 6 and 7 (BJ

1976.5.3-4) ; paratype 8 (Paris Muséum AIT 321). Stn 2 — paratypes 4 and 5 (BJ 1976.5.

1-2). Stn 4 — paratypes 1-3 (Paris Muséum Ali 322).

Remarks

The alTinities of this species appear ambivalent. It may be placed in the perionychin

genus Diporochaeta (Acanthodrilidae sensu Gates) if duplication of the characteristic pair

of nephridia in each segment is allowed in that genus. It may alternatively be assigned

to the megascolecin genus Celeiiella (Octochaetidae sensu Gates) on the grounds of tubular

prostates, calciferous (?) lamellae in XII1-XV, and an albeit simple meronephric condition

with réplication of the preseptal nephrostomes. Celeriella is, however, a homogeneous

genus especially with regard to the form of the prostates which extend through many

segments and hâve the muscular duct extending through two or more segments, necessi-

tating prolongation of the vas deferens well behind XV1I1. It seems préférable, therefore,

to associate this species with Diporochaeta (which absorbs ail Indian species currently in

Plutellus) regarding the duplication of the nephridia as a purely spécifie character of D. dor-

sochaeta. A similar réplication occurs in the typically holonephric Australian genus Diplo-

trema. The validity of associating this species with Diporochaeta is indicated by its close

482

B. G. M. JAMIESON

similarity to the purely holonephric Diporochaeta (= Plutellus) campsiaulus, also from

Palni Hills. Similarities of D. campsiaulus include the epilobous prostomium ; first dorsal

pore in 5/6 ; termination of the clitellum in XIII and XVII ; location of male and sperma-

thecal pores in b lines ; the general form of the spermathecae and, most striking, the un-

usually dorsal displacement of the dorsal setae. Significant différences of D. dorsochaeta

from D. campsiaulus are, however, the fact that dd is always less than cd (in the latter

species changing from 3/4 cd in the midbody to 2.5 cd in the forebody), the presence of

penial setae and the duplication of the nephridia.

Diporochaeta macrochaeta (Stephenson, 1925)

D. macrochaeta simpliciseta suhsp. nov.

(Fig. 1 A ; 5 H ; 6 H ; Table I)

Length 80 + mm, width (midclitellum) 1.7 mm, 143 + segments (H ; ail 7 specimens

are posterior amputées). Prostomium proepilobous small, in ail specimens withdrawn so

as not to be dorsally visible. First dorsal pore (minute) 10/11. Setae in eight regular

longitudinal rows as far as posterior amputation ; in XII aa : ab : bc : cd : dd : averaging

2.8 : 1.0 : 3.0 : 2.5 : 12.9 ; dd : u = 0.42-0.48 (mean 0.45, 5 specimens). Nephropores not

externally recognizable. Clitellum not visible but intersegments 16/17 and 17/18 dorsally

obscured in one specimen. Male pores well latéral of b lines of XVIII, relative to adjacent

segments, on distinct oval papillae winch are separated by a moat from a high, narrow

tumid wall wbich encloses both papillae, extends laterally to c lines, slightly overhangs

XIX and in the midline enters XVII, forming a médian tumescence in that segment ; a

médian extension of the wall passing forward from its posterior border between the papillae

to shortly anterior of the equator of XVIII. Female pores minute, shortly presetal in XIV,

almost contiguous medianly. Spermathecal pores small protubérances into 7/8 and 8/9

from VIII and IX, in a lines.

Thickest septa 7/8-12/13, strongly thickened, with 7/8-9/10 the strongest ; ail of these

funnel-shaped. Dorsal blood vessel single, continuous onto the pharynx ; last hearts in

XIII ; only those in X-XIII latero-oesophageal, each with a connective from dorsal and

supra-oesophageal vessel ; supra-oesophageal in IX-XI as wide as the dorsal vessel, tra-

ceable but narrow in at least VIII and XII and XIII ; suhneural vessel absent. Gizzard

in V, very large, strongly muscular and broadly fusiform, extending into IX by displace¬

ment of septa. Oesophagus in XII-XVI darkly vascularized and segmentally swollen ;

internally rugose, but extramural calciferous glands absent. Intestine commencing in XIX ;

muscular thickening and caeca absent ; a deep laminate dorsal typblosole starting almost

imperceptibly in XXII; intestine thicker wall in XXIV postenorly. Nephridia in VI,

VII and VIII (no nephridia seen anterior to this) forming increasingly large tufts lying

dorsolaterally on the oesophagus and sending each a very long, conspicuous (composite ?)

duct which enters the parietes at the anterior margin of the preceding segment in cd.

Succeeding nephridia a small exonephric (meronephric ?) tuft on each side ; no funnels

demonstrable ; ducts entering the parietes presetally in cd ; caudally with a pair of trans-

versly elongate nephridia, each with a preseptal funnel, on each side which appear to

MEGASCOLECIDAE FROM THE PALNI HILLS

483

Fig. 1. — Génital fields. A, Diporochaela macrochaeta simpliciseta subsp. nov. (holotype) ; B, Troyia

gundarshola gen. et sp. nov. (holotype).

Abbrevialions in figs. 1-6 : Ç, female pore ; g.m., aocessory génital marking ; (J, male pore ; p.s.,

penial seta ; pr.cl., prostate duct ; pr. g., glandular part of prostate ; sem. gr., séminal groove ; sp.p.,

spermathecal pore ; t.pub., tubercula pubertatis ; v.d., vas deferens. Roman numerals are segment

numbers. Clitellum shaded. Ail by caméra lucida.

be holonephridia though each has several small spiral loops medianly, the duct long and

very slender, entering the parietes presetally in cd. Metandric ; testes and iridescent

sperm funnels in XI ; testis-sacs absent ; large racemose séminal vesicles in XII. Ovaries

with several attenuated strings of large oocytes and funnels, in XIII ; ovisacs présent (?).

450, 3

484

B. G. M. JAMIESON

Prostates tubular, much eoiled in XVII1 and XIX, the long convoluted muscular duct

confined to XVIII and receiving the vas déférons at its entai third. Penial setae several

hair like, terminally wavy, without v isible ornamentation, the tip a fine point sometimes

bent ; several in a follicle ; a mature seta C.66 mm long, 3 pin wide at midlength. Sperma-

thecae two pairs, in VIII and IX, eaeh with elongate digitiform more or less tortuous

ductless ampulla, and similar but much smaller and shorter iridescent diverticulum (H, PI).

Materiai, examined : Stn 1 — holotype (Paris Muséum AI I 323), paratvpes 1-4 (Paris Muséum

AIT 324-327); paratypes 5 and 6 (BJ 1976.5.5-6).

Remarks

Diporochaeta macrochaeta was retained, with réservations, in Plutellus by Gates (1972).

The necessitv of hroadening the définition of Diporochaeta to include lumbricine species

with the perichaetine type-species (aide Jamieson, 1974a, 1976a) and restriction of Plu¬

tellus has here required inclusion, at least provisionally, of Oriental plutelli in Diporochaeta

but the generic status of D. macrochaeta is especiallv questionable. The new taxon is

included as a subspecies in D. macrochaeta , the nominate subspeeies of which is from the

Anamalai Hills, but it is possible that its departures from Stepuenso.n’s description indicate

spécifie distinction. Différences of D. macrochaeta macrochaeta include the following : male

pores are in ah on minute papillae in the absence of a surrounding tumescent wall ; location

of the gizzard mainly in VI (a questionable différence) ; origin of the intestine in XVIII ;

flattening and terminal notehing of the penial setae ; and the greater length of the sperma-

thecal diverticulum relative to tlie ampulla.

Tribe MEGASCOLECIN1 Jamieson, 1971a

Genus CELERIELLA Gates, 1958. Emend.

Small to moderate sized earthworms with approximately 100 segments. Prostomium

epilobous. First dorsal pore 4/5 or 5/6. Setae 12 to approximately 40 per segment,

commencing on II ; with significant middorsal and midventral gaps ; none between the

male pores. Clitellum annular, anterior to the segment bearing the male pores, occluding

intersegments and dorsal pores. Pores of the vasa deferentia and a single pair of tubular

prostates 1 pair on XVII1. Female pores a minute pair, anteromedian of setae a of XIV.

Spermathecal pores minute, 2 pairs in 7/8 and 8/9 or 1 pair in 7/8.

Some preclitellar septa thickened. Dorsal blood vessel single, continuous onto the

pharynx ; last hearts in XII ; those in X-XII latero-oesophageal ; supra-oesophageal limited

to the vicinity of these hearts ; subneural blood vessel absent. Gizzard large in VI. (Jeso-

phagus with high calciferous (?) lamellae in XIII and XIV but no extramural glands or

with a pair of discrète extramural pouches in each of XIII and XIV of which the anterior

pair sometimes communicates with the oesophageal lumen via the posterior pair. Intestine

MEGASCOLECIDAE FROM THE PALNI HILLS

485

commencing in XVI or XVII ; muscular thickening, caeca and typhlosole absent. Mero-

nephric ; in the oesophageal région with exonephric and at least in one species enteronephric

(pharyngeal) tufts followed by astomate, avesiculate, exonephric micromeronephridia. In

the intestinal région, caudally or throughout its length, with several to many preseptal

Fig. 2. — Génital fields. A, Celeriella quadripapillata (Stephenson, 1924) (specimen 1) ;

B, Diporochaeta dorsochaeta sp. nov. (holotype) ; C, l). dorsochaeta (paratype 4).

nephrostomal funnels on each side in each segment corrcsponding with postseptal (always

exonephric ?) avesiculate micromeronephridia ; megameronephridia absent. Holandric,

lestis-sacs absent ; séminal vesicles in XI and XII. Ovaries, with few to several eggstrings

in XIII, ovisacs présent, separate from the funnels in XIV. Prostates with 3 régions :

an anterior (ectal) wide muscular duct which extends through 2 or more segments, a slender

duct continuons with this and occupying a segment or less, and a very long, tortuous

slenderly tubular gland extending through several to more than 10 segments. Combined

vasa deferentia of a side joining the jmiction of gland and slender duct. Penial setae

486

B. G. M. JAMIESON

absent. Spermathecae with ovoid ampulla, short duct and a single clavate to digitiform

diverticulum.

Type species : Spenceriella duodecimalis Michaelsen, 1907.

Distribution : Palni Hills, S. India.

Remarks

Gates (1958) erected Celeriella for lndian species which had beeri placed in the Austra-

ûan genus Spenceriella. The latter genus has been revived by Jamieson (1972, 1974/j)

and its discreteness from Celeriella is confirmed. Important departures in the above

new définition from that of Gates are the démonstration of several to many preseptal

nephrostomal funnels (not merely one médian funnel) in caudal segments ; the inclusion

of species with extramural calciferous glands, and démonstration of union of the vasa

deferentia with the entai ends of the prostate ducts. The multiple caudal nephrostomes

confirm similarity which the lndian genus Priodoscolex (but not Priodochaeta) suggested

by Gates ; they also occur in Troyia and in the Australia généra Spenceriella and Oreos-

colex.

Table I. — Intersetal distances in segment XII as percentage of periphery.

ClRCUMFERENCE

(mm)

1). dorsochaeta

Holotype

15.6

8.1

13.9

17.1

9.2

15.3

13.9

6.9

3.5

Range from

12.0

5.7

11.1

6.9

14.5

13.0

6.6

15.9

9.3

17.5

20.5

13.2

18.1

15.9

10.2

Mean of 8

14.4

7.7

13.2

16.7

9.4

16.2

14.3

8.1

X intervals/ab

1). rnacrochaeta

simpliciseta

1.9

1.0

1.7

2.1

1.2

2.1

1.8

1.0

Holotype

8.4

3.6

10.8

7.8

48.2

7.8

9.6

3.0

6.6

Range from

8.4

3.0

8.9

7.5

41.6

7.3

9.6

3.3

11.2

3.9

12.0

13.2

48.2

8.9

11.2

3.9

Mean of 5

9.7

3.5

10.5

9.1

45.2

8.1

10.3

3.6

X intervals/ab

2.8

1.0

3.0

2.6

12.9

2.3

2.9

1.0

Lampito bouchei

Holotype

8.5

4.3

7.6

7.0

56.7

5.8

6.4

3.7

13.1

Paratype

10.0

4.3

7.7

6.2

53.5

7.3

7.1

3.9

11.2

Mean

9.3

4.3

7.7

6.6

55.1

6.6

6.8

3.8

interval/ah

2.3

1.0

1.8

1.6

13.6

1.5

1.6

0.9

T. gundarshnla

Holotype

7.5

3.8

9.7

5.9

50.7

8.0

10.7

3.8

14.9

interval/ab

2.0

1.0

2.6

1.6

13.3

2.1

2,8

1.0

MEGASCOLECIDAE FROM THE PALNI HILLS

487

Kby to species of Celeriella

1 .

2 .

6 .

Setae paired, 10-12 per segment, in at Ieast the forebody. 2

Setae not paired, 40-50 per segment. 6

2 pairs of spermathecae. 3

1 pair of spermathecae. 4

Spermathecal pores in the vicinity of a lines. Spermathecal diverticulum approximately half

the length of the ampulla. Longitudinal tubercula pubertatis each with a longitudinal row of

minute orifices in 14 XVI1-14 XIX. C. punctata sp. nov.

Spermathecal pores in bc lines. Spermathecal diverticulum more than twice the length of

ampulla and duct. No tubercula pubertatis. C. ditheca (Stephenson, 1924)

Spermathecal pores in or just latéral to b lines. 5

Spermathecal pores in or close to d lines. C. quadripapillata (Stephenson, 1924)

Setae in 12 regular longitudinal rows throughout the body.

C. regularis (Stephenson, 1924)

Only a and b setae in regular rows throughout ; in the hindbody other rows irregular and

number increased to 14-17 per segment. C. duodecimalis (Michaelsen, 1907)

Male papillae medianly contiguous ; male pores lying in a transverse midventral groove but

not at the anterior ends of longitudinal grooves ; ectal muscular duct of prostate considerably

shorter than gland. C. kempi (Stephenson, 1924)

Male papillae not medianly contiguous ; male pores not in a transverse midventral groove

but each at the anterior end of a comma-shaped séminal groove ; ectal muscular duct of prostate

at least twice as wide as the gland. C. bursata sp. nov.

Celeriella bursata sp. nov.

(Fig. 2 A ; 4 A ; 6 A ; Table 2)

Length 82 mm, width (midclitellum) 2.6 mm, segments 110. Prostomium epilobous %,

open, narrow and parallel-sided. First dorsal pore 5/6. Setae 40 in XII, 42 in XX ;

in XII aa : ab : zy : zz = 3.0 : 1.0 : 1.0 : 1.5 : 5.2 ; no setae between the male pores. Nephro-

pores not externally visible. Clitellum not developed. Male pores in setal lines 4 on a

pair of conspicuous hemispheroidal papillae in XVIII, each pore at the anterior end of a

broad séminal groove which runs posteromedianly to the posterior edge of the porophore

and then constinues on a tumid extension to the posterior margin of XVIII, the porophore

plus extension having the form of a comma. Female pores minute, shortly anteromedian

of setae a of XIV. Spermathecal pores minute, concealed in intersegmental furrows 7/8

and 8/9, in ab ; those in 8/9 0.44 mm apart.

Thiekest septa 10/11 and 11/12, very strong. Dorsal blood vessel single ; continuons

onto the pharynx ; last hearts in XII ; only those in X-XII latero-oesophageal, each with

connective from dorsal and supra-oesophageal vessel ; supra-oesophageal well developed,

in X-XIV ; subneural vessel absent. Gizzard in VI, very large and strongly muscular

and fusiform, extending into X by displacement of septa. Oesophagus with true extra¬

mural calciferous glands, a pair in each of XIII and XIV ; each gland a large latéral pouch

broadly attached to the oesophagus, without. narrow connection, but its lumen divided

488

B. G. M. JAMIESON

by numerous radial longitudinal lamellae which run from the outer wall and join the external

wall of the oesophagus, the mode of opening into the oesophagus not determined. Intestine

commencing in XVII ; muscular thickening, caeca and typhlosole absent. Nephridia

paired tufts in III, IV and V enlarging posteriorly to large in V, with multiple discrète

ducts discharging (ail ?) exonephrically ; a few small nephridia forming a transverse row

anteriorly in 111 ; the équivalents of the tufts in VI and VII becomming diffuse laterallv

and a pariétal transverse band of exonephric astomate micromeronephridia developed by

VIII. By XIII these nephridia are much smaller and are scattered on the parietes. In

the anterior intestinal région in the order of 10 more elongate but otherwise similar nephri¬

dia on each side in each segment. Caudally with approximately 15 long stalked preseptal

nepbrostomal funnels on each side on the posterior septum of each segment, the nephri-

dial hodies densely clothing the posterior face of the septum ; some if not ail of these

nephridia exonephric ; no megameronephridia. Holandric, funnels iridescent in X and

XI; testis-sacs absent; much subdivided racemose séminal vesicles in XI and XII.

Ovaries small, with several strings of large oocytes, and funnels in XIII ; ovisacs

absent. Prostates tubular, immensely long, the glandular portion very tortuous and

extending from XXII to XXXIII, passing anteriorly into a narrow winding tube with

spermatozoal (?) iridescence (intermediate tube) which runs through XXI to posterior

XX (receiving the vas deferens posteriorly), becomes strongly muscular and wider al

its anterior end and passes into an unusually wide very muscular duct (bursa) extending

from it to the male pore equatorialy in XVIII, narrowing shortly liefore the pore. Vas

deferens traced from XVII to the ectal end of the widest part of the terminal

bursa but emerging again to run down the intermediate tube and apparently joining the

entai end of the latter. Spermathecae 2 pairs in VIII and IX, each with a large ovoid

ampulla and slightly longer digitiform diverticulum, more than lialf as wide ; free extremity

of diverticulum slightly swollen and with spermatozoal iridescence.

Matériau examined : Stn 1 — holotype (Paris Muséum AH 328).

Remarks

Celeriella bursata is morphologically close to C. kempi (Stephenson, 1924) the only

other species of the genus with more than 12 setae in anterior segments. The latter species

is known only from a single apparently immature and imperfectly characterized specimen.

Distinction of C. bursata from it is therefore not as positive as would be désirable. Diffé¬

rences from C. kempi as described by Stephenson are as follows. The male papillae are

not medianly contiguous ; the male pores do not lie in a transverse midventral groove ;

a longitudinal séminal groove is présent, extending onto a posterior prolongation of each

male porophore ; although the terminal prostate duct in C. kempi is “ shiny, and conside-

rably stouter ” than the remainder, the enormous muscular thickening of the ectal duct

in C. bursata is clearly of a different order, rarely attained in megascolecines, meriting appli¬

cation of the terni bursa. Only one pair of structures questionably recognizable as sperma¬

thecae has been seen (in IX) in C. kempi but it would be prématuré to State that the two

taxa difîer with respect to their spermathecae. The possibility nevertheless remains

MEGASCOLECIDAE FROM THE PALNI HILLS

489

Fig. 3. — Génital fiolils.

A, Celeriella bursala sp. nov. (holotype) ; B, Celeriella punctata sp. nov. (lmlotype).

that, diseovery and examination of further material of C. kempi from the type-locality,

Mariai) Shola in the Palni Hills, may narrowthese distinctions and may necessitate regarding

C. bursata as its junior synonym.

Celeriella punctata sp. nov.

(Fig. 3 B ; 4 B ; 6 B ; Table TI)

Length 80, 75, 84 mm, vvidth (midclitellum) 2.1, 2.0, 1.8 mm, segments 137, 127, 130

(holotype and paratypes 1 and 2 ; paratype 3 is a posterior amputee). Prostomium epilo-

bous 1/4, broad closed. First dorsal pore 5/6. Setae 12 per segment throughout, in 3

490

B. G. M. JAMIESON

regular pairs on each side, the ventral pair ( ab ) closely paired, the next pair (cd) moderatelv

closely, the third pair (zy) widely paired ; zz and aa wide interruptions tbroughout, zz

especially wide anteriorly but little wider tban zy caudally ; in XII aa : ab : bc : zy : zz

averaging 2.4 : 1.0 : 1.5 : 1.8 : 6.0 (H, Pl-3). Nephropores not externally recognizable.

Clitellum annular, % XIII-XVII. intersegments, dorsal pores and setae occluded. Male

pores a pair of minute orifices equatorialy in XVIII in a lines, each pore on an indistinct

papilla which is part of a longitudinal tumescence (tuberculum pubertatis) which extends

from the equator of XVII almost to the equator of XIX ; each tuberculum bearing a longi¬

tudinal row of closely set dark points in single file in ab extending throughout its length,

excepting the small male papilla ; 11 points between the setal arc of XVII and the male

papilla, and 4 points behind it, on the right tuberculum (H, Pl-2) ; no longitudinal rows

of points visible in P3. These points shown in longitudinal sections (P4) to be each the

orifice of a hemispheroidal thickly muscular organ. Female pores minute, anteromedian

to setae a of XIV. Spermathecal pores 2 pairs, in 7/8 and 8/9, on minute papillae imrne-

diately latéral of a lines.

Thickest septa 6/7-8/9, moderately strongly thickened. Dorsal blood vessel single,

continuous onto the pharynx ; last hearts in XII ; only those in X-XII latero-oesophageal,

each receiving a connective from the supra-oesophageal vessel and an exceedingly slender

one from the dorsal vessel ; supra-oesophageal in X-XII, well developed ; subneural vessel

absent. Gizzard in VI, large and strongly muscular, cylindrical, extending to % IX by

backward deflection of septa. Oesophagus with 2 pairs of large, very discrète, ovoid

calciferous glands, in XIII and XIV, the larger anterior pair, though appearing discrète,

passes at the posterior end into the glands of XIV of which tbey are merely an expansion

anterior to septum 13/14 ; the glands of XIV attached by their ventral pôles to the oeso¬

phagus posteriorly in the segment ; the lumen with numerous high vascular laminae which

almost occlude the lumen of the gland but do not fuse across it. Intestine commencing

with abrupt expansion in XVI ; muscular thickening, caeca and typhlosole absent. Nephri-

dia : two pairs of very large tufts in V and VI (the latter at the anterior limit of the gizzard)

send composite duct.s forward to join the wall of the pharynx ; there is also a pair of small

tufts posteriorly in each of VI and VII, diseharging exonephrically in their respective

segments and il is likely, therelore, that the large anterior tufts in VI belong to V, or even

further anteriorly, septum 5/6 being unrecognizable ; diffuse exonephric tufts, scarcely

sufïiciently compact to be considered tufts, in VIII and IX ; rudimentary tufts and a few

pariétal meronephridia in X and XI ; sparse astomate exonephric, avesiculate pariétal mero-

nephridia in XII posteriorly. Caudally with several minute preseptal nephrostomal

funnels on each side in each segment ; some at least of the nephridia exonephric, a trans¬

verse wad of nephridial tubules near the anterior septum and adhèrent to tbe dorsolateral

aspect of the intestine is apparently also exonephric by several ducts though enteronephry

cannot be considered disproved. Holandric : iridescent sperm funnels in X and XI ; testis-

sacs absent ; much-divided racemose séminal vesicles in XI and XII. Üvaries, with few

conjoined strings of large oocytes, and funnels in XIII ; large ovisacs in XIV dorsal to

the oesophagus, not connected with the oviductal funnels. Prostates slenderly tubular,

very long, the glandular portion tortuous, though linear, extending tbrough XIX to XXVII,

passing anteriorly at % XX, into a narrow sinuous duct which continues into XIX, trans-

forming in this into a wide muscular duct, also sinuous, which discharges equatorialy in

MEGASCOLECJDAE FROM THE PALNI HILLS

491

XVIII in a lines ; vas deferens joining the junction of gland and duct. The longitudinal

sériés of hemispheroidal glossy swellings of the tubercula pubatatis and the male pores

overlain by laterally radiating muscle bands. Spermathecae 2 pairs, in VIII and IX,

the posterior pair significantly the larger ; each with ovoid sometimes apically lobed ampulla

and slender tortuous duct which is joined at midlength by a médian clavate inseminated

diverticulum which is almost half the length of the ampulla.

Material examined : Stn 5 — holotype (Paris Muséum AII 329), paratypes 1-2 (Paris Muséum

AH 330-331) ; paratype 3 and (sections) 4 (BJ 1976.5.7-8).

Remarks

The only other quadrithecal species of Celeriella, at least with 12 setae per segment,

is the inappropriately named C. ditheca (Stephenson, 1924). This is again, very inade-

quately described from a single specimen but the new material here named C. punctata,

differs from Stephenson’s description of C. ditheca in the following respects. Spermatheeal

pores are in the vicinity of a lines (not in bc ) ; the spermatheeal diverticulum is only about

half the length of the ampulla (not approximately twice the length of ampulla 4- duct) ;

prostates end in XXVII (not XXXIII or XXXIV) ; calciferous glands are better developed

in XIV than in XIII and dorsal pores commence in 5/6 (not 4/5). The remarkable tubercula

pubertatis with their longitudinal rows of minute dark orifices hâve not been reported

for C. ditheca.

Celeriella quadripapillata Stephenson, 1924

(Fig. 2 A ; 5 C ; 6 C ; Table II)

Length 68, 62, 84 mm, width (midclitellum) 1.6, 1.4, 1.6 mm, 91, 98, 105 segments

(specimens 1-3). Prostomium epilobous 34; small, open, parallel sided. First dorsal pore

4/5 (1,2). Setae in 12 longitudinal rows throughout, though 1 of each ventralmost pair

is absent in XII in specimens 1 and 5 ; in XII aa : ab : bc : zy : zz averaging 2.2 : 1.0 : 1.3 :

1.2 : 2.0 (5 specimens) ; aa conspicuously wider, zz hardly appreciably wider than adjacent

intervals throughout. Nephropores not externally recognizable. Clitellum % XIII-

XVI, weakly developed, intersegmental furrows slightly obscured. Male génital field a

large low but distinct circular tumescence fringed by concentric rings centred on ab and

filling the posterior three-fourths of segment XVIII ; each tumescence bearing a pair of

small papillae side by side and almost pinched together, the latéral one in b line, the médian

one in ab relative to adjacent segments ; the male pore situated between the papillae and

almost concealed by them. The body ventrally flattened in the vicinity of the tumescences

and depressed between them so that the tumescences occupy longitudinal ridges. Female

pores shortly anteromedian of setae a in XIV. Spermatheeal pores i pair, in 7/8, in d.

lines, minute with slight peripheral tumescence.

Thickest septa 6/7 and 7/8, moderately strongly thickened. Dorsal blood vessel single,

continuons onto the pharynx ; last hearts in XII, oïdy those in X-X1I latero-oesophageal,

each with a connective from the dorsal and supra-oesophageal vessel ; supra-oesophageal

well developed, in at least 34 VIII-XII ; subneural vessel absent. Gizzard in VI, mode-

492

B. G. M. JAMIESON

rately large, strorigly muscular, globose, extending to }4 VIII by backward deflection of

septa. Oesophagus dilated in each of XIII and XIV and with numerous free bigh internai

radial lamellae extending into its lumen but with no separate extramural calciferous glands.

Intestine commencing in XVI but not considerably widening until XVII ; muscular

thickening and eaeca absent ; a very rudimentary dorsal ridge présent from XXIII does

not warrant récognition of a typhlosole. Nephridia : a very large pair of tufts in each

of VI and VII sending a composite duct into V and VI, respectively, on each side which

appears to be exonephric. Astomate exonephric micromeronepbridia in A'111-X several

scattered on the parietes ; by XI forming a sparse transverse band, the band becomming

more dense with some médian ventral aggregation by XIV in which some nephridia appear

to hâve intrasegmental nephrostomes ; caudally with several preseptal nephrostomes on

each side but no discernible enteronephry. Holandric ; iridescent funnels in X and XI ;

testis-sacs absent ; mucb-divided racemose séminal vesicles in XI and XII. Ovaries

small with several fused strings of large oocytes, and funnels in XIII ; ovisacs absent.

Prostates slenderly tubular, immensely long, the glandular portion tortuous and extending

through XXII-XXVI, passing anteriorly into a narrow wdnding tube which runs through

XXI (receiving the vas deferens posteriorly) and in XX to the pore in XVIII forms a very

wide, glossy muscular terminal bursa. Spermathecae 1 pair, in VIII, each with ovoid-

reniform ampulla and narrowly tapering indistinctly demarcated duct which enters the

body wall in common with a long, twisted, inseminated, digitiform diverticulum which

is longer than ampulla plus duct.

Materiai. examined : Stn 2 — 1 (BJ 1976.5.9). Stn 3 — 4 (Paris Muséum AU 332-AH

335) ; l (B.I 1976.5.10).

Tvpe-Locality : Kodaikanal, Palni ITills.

Genus TROYIA nov.

Setae 8 per segment, commencing on II, widely paired. Clitellum annular, anterior

to the segment bearing the male pores. Pores of the vasa deferentia and a single pair

of tubuloracemose prostates 1 pair on XVIII. Female pores a minute pair, anteromedian

of setae a of XIV. Spermathecal pores a pair at the anterior margin of VIII.

Some preclitellar septa thickened. Dorsal blood vessel single, continuous onto the

pharynx ; last hearts in XII ; those of X-XII latero-oesophageal ; supra-oesophageal

limited to the vicinity of these hearts ; subneural vessel absent. Gizzard large, in VI

or VII (?). 3 pairs of discrète extramural calciferous glands in XIII, XIV and XV, ven-

trallv attached to the oesophagus by narrow connectives but virtually sessile, their walls

with numerous free radial laminae. Intestine commencing in XVI ; muscular thickening,

caeca and typhlosole absent. Meronephric ; enteronephric tufts in V ; exonephric (?)

tufts in VI-XIII ; caudally with numerous preseptal nephrostomal funnels on each side

in each segment corresponding with postseptal exonephric avesiculate micromeronephridia ;

megameronephridia absent. Holandric ; testis-sacs absent ; séminal vesicles in XI and XI1.

Ovaries with many egg strings in XIII ; ovisacs présent, separate from the funnels, in XIV.

MEGASCOLECIDAE FROM THE PALNI HILLS

493

Table il. — Intersetal distances in segment XII as percentage of periphery.

a a

a b

z y

ClRCUMFER K NC

(mm)

('. hursata

Holotype

4.0

1.3

1.4

2.0

6.7

9.1

interval/ab

3.0

1.0

1.5

5.3

C. punctata

Holotype

11.3

5.0

5.4

10.1

31.5

3.7

Range from

7.9

3.7

5.4

5.5

23.2

12.5

5.0

9.0

10.1

31.5

Mean of 4

10.1

4.3

6.6

7.8

25.9

interval/ab

2.4

1.0

1.5

1.8

6.0

C. quadripapillala

Holotype

13.4

6.7

7.8

6.7

14.2

5.4

Range from

12.1

5.6

7.5

6.1

9.5

15.6

7.1

9.8

11.7

17.1

Mean of 5

13.9

6.4

8.3

7.6

12.9

interval/ab

2.2

1.0

1.3

1.2

2.0

L. sylvicola

Range from

9.2

3.9

4.3

4.8

28.3

12.4

5.2

7.2

(>.7

38.8

Mean of 5

10.6

4.3

5.9

5.5

33.6

interval/ab

2.5

1.0

1.4

1.3

7.9

Prostates witli 3 régions, an anterior (ectal) wide muscular duct which extends through

more tlian 1 segment, a short slender duct continuons with this and a linear tubuloracemose

gland extending through several segments. Combined vasa deferentia of a side joining

the junction of gland and slender duct. Penial setae absent. Spermathecae with single,

clavate diverticulum.

Type species : Troyia gundarshola sp. nov. (Monotypic).

Distribution : Palni Hills, S. India.

Remarks

Troyia is very closely related to Celeriella of which it is here regarded as the apomorph

sister-group, its apomorph characters relative to Celeriella being the third pair of calci-

ferous glands, in XV, and the tubuloracemose rather than tubular form of the prostates.

If presence of calciferous glands in XIII-XV were treated as a spécifie rather than a generic

character, T. gundarshola would probably bave to be placed in Celeriella.

494

B. G. M. JAMIESON

Troyia gundarshola sp. nov.

(Fig. 1 B ; 5 D ; 6 D ; Table I)

Length 273 mm, width (midclitellum) 7.0 mm, 297 segments. Prostomium proepilo-

bous but impinging on about 1/3 of the peristomium (H) or epilobous 2/3 (PI). First dorsal

pore 5/6. Setae in 8 longitudinal rows, a developing slight sporadic irregularity posteriorly

but d and then c and b becoming very irregular ; in XII (H) aa : ab : bc : cd : dd — 2.0 :

1.0 : 2.7 : 1.9 : 13.3 ; dd : u — 0.51. Nephropores not externally visible. Clitellum

annular, strongly tumid, 1/3 XIII-XVII, intersegmental furrows partially obscured ventrally.

Male pores in ab, near a. of XVIII, relative to adjacent segments, each with wide tumid

bps which constitute a papilla ; a short inconspicuous groove extending from each pore

onto the anterior lip may be a séminal groove ; the two papillae medianly contiguous.

Female pores minute, anteromedian to setae a of XIV. Spermatbecal pores a pair of

indistinct slits near the anterior margin of VIII slightly médian of a lines, in a common

tumid area, with the appearance of closed lips, the anterior and posterior lip filling the

corresponding halves of segments Vil and VIII.

Thickest septa 8/9-12/13, strongly thickened, with 9/10-11/12 the strongest ; ail of

these funnel shaped. Dorsal blood vessel single, continuons onto the pharynx ; last hearts

in XII ; only those in X-XII latero-oesopbageal, each with a connective from dorsal and

supra-oesophageal vessel (H, P). Supra-oesophageal in posterior IX to anterior XIII,

well developed. Subneural vessel absent. Gizzard in VII, i.e. abutting on septum 7/8

posteriorly without intervention of other septa (H, possibly anterior to the extremely

attenuated septum 6/7 in P) ; large, strongly muscular, broadly cylindrical with anterior

rim ; extending into X by backward displacement of septa. Oesoph agus with 3 pairs of large

very discrète calciferous glands, a pair in each of XIII, XIV and XV (II, P) ; each gland

ovoid, with the longest axis approximately vertical, and with a very narrow attacbment

at its ventromedian extreme to the oesophagus without intervention of an appréciable

stalk ; a transverse section of the gland reveals numerous closely packed radial septa or

laminae which almost occlude the lumen but do not join across it ; the laminae mostly

free axially but some fused with adjacent laminae. Intestine commencing in XVI ; muscular

thickening, caeca and typhlosole ahsent (H, PI). Nephridia : very small paired exonephric

tufts in (II ?), 111 and IV ; large paired astomate, enteronephric tufts in V sending long

composite ducts forwards to the anterior limit of the pharynx ; paired tufts in VI-XIII

becoming smaller and more diffuse, (ail ?) exonephric. Integumentarv nephridia in XIV

XVII very numerous minute astomate micromeronephridia covering the entire parietes.

In XVIII posteriorly each segment with moderately numerous but sparsely distributed

astomate exonephric micromeronephridia on each side. Caudally with a pariétal band

of (exonephric ?) meronephridia and numerous minute (enteronephric ?) micromerone¬

phridia on the posterior face of each septum corresponding with small, long-stalked presep-

tal funnels ; no megameronephridia présent. Holandric ; sperm funnels iridescent in X

and XI ; testis-sacs ahsent ; racemose much divided séminal vesicles in XI and XII (II, P).

Ovaries with many strings of large oocytes, and funnels,' in XIII ; large structures with

many moniliform projections in XIV appear to be ovisacs (H, P). Prostates tuhulora-

MEGASCOLECIDAE FROM THE PALNI HILLS

495

cemose, a pair in XVIII-XXV, the glandular portion, in XX-XXV, depressed and alter-

nately incised so that it is still referable to a tortuous depressed tubular form ; muscular

octal duct narrow and coiled for a short portion near the gland but of moderate and uniform

width for most of its length ; receiving the vas deferens at junction with the gland (II) ;

in the paratype the prostate duct extends through XVIII-XX ; the glandular portion is

thickly strap-like, in posterior XX-XXV. Penial setae absent. Spermathecae in VIII,

basicallv a pair but that on the left with anipulla and diverticulum completely duplicated,

the two spermathecae smaller than the single right spermatheca which bas large ovoid

slenderly tapering but virtually duetless ampulla and a single médian clavate longer diver¬

ticulum which joins it hasally.

Materiai, examined : Stn 5 — holotype (Paris Muséum AH 336). Stn 4 — paratvpe (lîJ

1976.5.121.

Il EM AI! KS

Location of extramural calciferous glands in XIII-XV distinguishes T. gundarshola

front ail other Indian megascolecids.

Getms LAMPITO Kinl >erg, 1866

Dorsal pores présent. Setae 8 to numerous per segment. Clitellum annular. Pores

of the vasa deferentia and a single pair of racemose or tuhuloracemose (strap-shaped)

prostates 1 pair on XVIII. Fentale pores paired on XIV. Spermathecae 2 or 3 pairs,

pores intrasegmental, or at the anterior intersegmental furrows of VII, VIII and IX.

Vascular System with unpaired dorsal, ventral and supra-oesophageal trunks but no

suhneural, paired extra-oesophageals médian to the hearts, paired lateroparietal trunks

from the anal région connecting in XIII with extra-oesophageal and supra-oesophageal

trunks; last hearts in XIII; those in X-XIII (sometimes only XII and XIII) latero-

oesophageal. Gizzard in V. Calciferous tissue in longitudinally placed, more or less

lamelliform ridges on the inner wall of the oesophagus in X-XIII. Intestine commencing

in XV-XVI, typhlosole présent or [L. bouchei) absent ; muscular thiekening and caeca

absent. Meronephric ; ail nephridia avesiculate, paired clusters (tufts) of astomate micro-

meronephric tubules on anterior faces of septa in V-(XI ?) XIII, XIV, ducts from some

clusters opening into the pharynx ; numerous astomate v-shaped micromeronephridia per

segment on the pariet.es from (XIII or XIV ?) XV posteriorly, in transverse hands ; the

médian nephridium on each side enlarged as an enteronephric megameronephridium with

preseptal funnel from XX or XXI posteriorly ducts passing to the roof of the intestine

and (alwavs ?) joining a médian longitudinal supra-intestinal canal from which a pair

of ductules opens into the gut in each segment. Holandric or metandric ; gymnorchous ;

séminal vesicles in XI and XII, or XII only. Prostates tuhuloracemose or racemose ;

vasa deferentia joining the entai ends of the prostate ducts. Penial setae présent or absent.

MEGASCOLECIDAE FROM THE PALNI HILLS

497

Key to species of Lampito

1 .

■)

6 .

Sperm funnels in X and XI .

Sperm funnels in XI only.

Spermathecal pores 2 pairs ; setae 8 per segment.

Spermathecal pores 3 pairs ; setae more than 8 per segment. L. mauritii Kinberg,

Male pores in a single field, typldosole bilid ventrally. L. marianae (Stephenson,

Male pores not in a single field, typblosole not bifid ventrally.

Spermathecal pores with same intrasegmental location in VIII as in IX.

Spermathecal pores postsetal in Y11 1, presetal in IX. L. kempi Gates,

Spermathecal pores presetal. L. vilpattiemis (Michaelsen,

Spermathecal pores not presetal.

Spermathecal pores équatorial. L. kumüiensis (Aiver,

Spermathecal pores postequatorial. L. sylvicola (Michaelsen,

Séminal vesicles in XI and XII. A very low typhlosole présent.

L. p:ilnicnsis (Stephenson,

Séminal vesicles in XII only. Typhlosole absent. L. bouchei sp.

18 ( 11 )

1924)

1938

1997)

1929)

1907)

1924)

nov.

Lampito bouchei sp. nov.

(Fig. 4 B ; 5 G ; 6 J ; Table I)

Length 80 +, 195 mm (H, posterior amputee ; P), width (midelitellum) 5.2, 3.8 mm,

304 segments (P). Prostomium proepilobous ?, in both specimens withdrawn and not

visible. First dorsal pore (minute) 10/11. Setae in 8 regular longitudinal rows throughout :

in XII aa : ab : bc : cd : dcl averaging 2.3 : 1.0 : 1.7 : 1.6 : 13.6 ; dd : u = 0.54 — 0.57 (2 spe¬

cimens). Nephropores not externally visible. Clitellum orange-brown, very strongly

tumid ; 1/3 XIII-1/3 XIX, intersegmental furrows obscured (H.) Male pores very slightly

latéral of b Unes of XVIII, relative to adjacent segments, on conspicuous medianly conjoined

hemispheroidal papillae which overhang tbe anterior and posterior limits of the segment

and are jointly circumscribed bv an indistinct tumid band which intrudes on XVII

and XIX (H, P). Female pores minute, anteromedian to setae a of XIV (P). Sperma¬

thecal pores small but conspicuous papillae in 7/8 and 8/9, in a Unes, the adjacent segments

puckered in their vicinity (H, P).

Thickest septa 5/6-12/13, strongly thickened, with 7/8-10/11 the strongest ; ail of

these funnel shaped. Dorsal blood vessel single, continuons onto the pharynx ; last heart.s

in XI11 ; those in X-XIII latero-oesophageal, each with a connective from dorsal and supra-

oesopbageal vessel (H, P). Supra-oesophageal % IX-XIII, well developed ; subneural

vessel absent. Gizzard in V, large, strongly muscular, globose but with anterior rim,

extending to IX by displacement of septa. Oesophagus in VIII-XIII with numerous

délicate high radial laminae, especially in XII-XII1, but these projecting into the general

oesophageal lumen and extramural calciferous glands absent. Intestine commencing

in XVI (P) ; muscular thickening, caeca and typhlosole absent. Nephridia : a pair of

498

B. G. M. JAMIESON

large tufts in V at the anterior limit of the gizzard sending a thick duct on each side to the

pharynx; a pair of large tufts in each of VI-1X send a composite duct forward on eacli

side traced to the junction of the anterior septum of each segment with the ventral parietes.

Smallers tufts in X and XI. By the posterior oesophageal région (XV) there are transverse

pariétal bands of bushy apparently astomate and exonephric meronephridia. From XX1

there are several small pariétal astomate exonephric micromeronephridia on each side in

transverse single fde but the median-most nephridium is enlarged as a megameronephri-

dium, with preseptal funnel, the body of the nephridium extending around the gut almost

Lo the dorsum, the long, narrow duct of each running on the anterior septum and entering

the intestine near the dorsal vessel ; this condition persisting to the caudal extremity but

the megameronephridium there becoming very slender, though still extensive laterallv ;

presence or absence of a longitudinal excretory duct not determinable. Holandric with

incipient metandry : large testes and iridescent funnels in XI with large racemose séminal

vesicles in XII ; in X with non-iridescent sperm funnels and no évident testes ; testis-sacs

absent. Ovaries, with many strings of large oocytes, and funnels in XIII ; large séminal

vesicle-like structures on the anterior septum of XIV may be ovisacs. Prostates, 1 pair

in XVIII and XIX, elongate racemose with secondary lobes which indicate internai bran-

ching of ducts ; ectal muscular duct forming a loop of moderate length and receiving the

vas deferens at its junction with the gland. Penial setae absent. Spermathecae 2 pairs,

in VIII and IX, each with elongate clavate ampulla fdling the segment longitudinally

and with 2 basal inseminated diverticula which may be simple, bifid, or trifid.

Material examined : Stn 1 — holotype (Paris Muséum AH 337) ; paratype (BJ 1976.5.13).

Remarks

Lampito bouchei is morphologically close to L. palniensis (Stephenson, 1924), the only

other species of Lampito with only 8 setae per segment throughout the body. It is here

separated with some hésitation from L. palniensis hecause it possesses the following dis¬

tinctions : well developed male porophores and an apparently different male génital lield ;

restriction of the séminal vesicles to XII ; total absence of a tvphlosole ; and presence of

latero-oesophageal hearts in X-XIII (not merely XII and XIII ?).

Lampito sylvicola Michaelsen, 1907

(Fig. 4 A ; 5 F ; 6 I ; Table II)

Length 198 mm, width (midclitellum) 4.2 mm, 239 segments (specimen 1 ; other spé¬

cimens are posterior amputées). Epilobous, c. % ? (1 specimen, withdrawn in other 4 spé¬

cimens). First dorsal pore 10/11. Setae 16 in XII, 30 in XX, 42 in a caudal segment ;

aa and zz conspicuous gaps excepting caudally where aa is no larger than other intersetal

distances ; in XII aa : ab : bc : zy : zz averaging 2.5 : 1.0 : 1.4 : 1.3 : 7.9 (specimens 1-5).

Nephropores not externally recognizable. Clitellum strongly tumid, 1/4 XII1-2/3 XVIII

(= 5

segments) ; intersegmental furrows and dorsal pores occluded ; setae retained.

MEGASCOLECIDAE FROM THE PALNI HILLS

499

/v.d

Fig. 5. — Prostate glands. A, Celeriella bursata sp. nov. (holotype) ; B, C. punclata sp. nov. (holotype) ;

C, C. quadripapillata (Stephenson, 1924) (specimen 1) ; D, Troyia gundarshola sp. nov. (holotype) ;

E, D. dorsochaeta sp. nov. (paratype 1) ; F, Lampito sylvicola Michaelsen, 1907 (specimen l) ; G, Lam-

pito bouchei sp. nov. (holotype) ; H, Diporochaeta macrOchaeta simpliciseta subsp. nov. (holotype).

Male pores in ah of XVIII (relative to adjacent setigerous segment) on minute papillae,

each papilla flanked by an ovoid tumescent génital marking ; the papilla and marking

lying in a deep oval dépréssion in a common oval strongly tumid pad which lills XVIII

longitudinally and extends between the two dépréssions. The latéral génital marking

not apparent in some specimens. Female pores probably paired, represented by a trans-

500

B. G. M. JAMIESON

Fig. 6. — A-J, Spermathecae. A, Celeriella bursata sp. nov. (holotype) ; B, C. punctata (holotype) ; C, C. qua-

dripapillata (Stephenson, 1924) (specimen 1) ; D, Troyia gundarshola sp. nov. (holotype) ; E-G, Dipo-

rochaeta dorsochaeta sp. nov. : Ë, holotype ; F, paratype 2 ; G, paratype 1 ; H, Diporochaeta macrochaeta

simpliciseta subsp. nov., right spermathecae of VIII and IX in situ ; I, Lampito sylvicola Michaelsen,

1907 (specimen 1) ; J, Lampito bouchei sp. nov. (holotype) ; K, Diporochaeta dorsochaeta sp. nov., left

penial setae (holotype).

verse slit presetally in XIV. Spermathecal pores 2 pairs of minute papillae at the posterior

border of the setal annulus in a lines in VIII and IX.

Thickest septa 6/7-9/10, strongly thickened and funnelshaped. Dorsal blood vessel

single, continuons onto the pharynx ; last hearts in XIII ; only those in X-XIII lotero-

oesophageal, each with a connective from the dorsal and supra-oesophageal vessel. Supra-

oesophageal in X-XIII, well developed only in XIII ; subneural vessel absent. Gizzard

in V, large, strongly muscular, fusiform extending to X by backward displacement of septa.

MEGASCOLECIDAE FROM THE PALNI HILLS

50]

Oesophagus lacking extramural calciferous glands. Intestine commencing in XVI ; mus-

cular thickening and caeea absent ; a low but distinct, broadly laminar sinuous dorsal

typhlosole commencing in XXVII. Nephridia : a pair of very large meronephric tufts

in V at the anterior limit of the gizzard send a composite duct forward, on each side, into II

in the vicinity of the buccal cavity into which it probably opens. Paired tufts in VI-XIV

diminish to virtually simple nephridia in XIV ; each sending a long (composite ?) duct

on each side to the ventral body wall at which it appears to fan out as a diaphanous mem-

branous structure ; no other nephridia présent in these segments. In XV-XIX there are

transverse bands of bushy, apparently astomate (exonephric ?) micromeronephridia at

the anterior border of each segment but in XX these are augmented by pariétal bands at

the posterior border and the médian nephridium is greatly enlarged, traversing the entire

width of the segment, as a megameronephridium ; a preseptal funnel is demonstrable for

the anterior-most and many subséquent megameronephridia ; a slender duct demonstrable

in the anterior intestinal région and caudally running from each megameronephridium to

the roof of the intestine beneath the dorsal vessel. Metandric ; testes and iridescent funnels

in XI and large racemose séminal vesicles in XII ; testis-sacs absent. Ovaries as flattened

lobes with a fringe of terminal oocytes, and funnels, in XIII ; lobulated structures in XIV

are presumably ovisacs. Prostates a pair, with large flattened glands in XVIII and XIX

which are superficially lobulated and incised but appear resolvable into an S-shape ; with

tortuous moderately long muscular ectal duct joined at the gland by the vas déférons.

Spermathecae 2 pairs, in VIII and IX, each with bulbous ampulla and narrowly tapering

sinuous duct ; 2 ellipsoidal inseminated diverticula joining the entai end of each duct, one

on each side ; the ducts in VIII and IX entering the body wall immediately behind setae a.

Material examined : Stn 5-2 (Paris Muséum Ail 338 (illustrated) — AH 339 ; 3 B J 1976.

5.11, 14, 15).

Type locality : Tiger Shola (near Kodaikanal), Palni Hills.

Remarks

The above description confirms location of both pairs of spermathecal pores behind

the setal arcs of tlieir respective segments, a location questioned by Gates (1938). The

génital marking latéral to the male pore was absent from the single type-specimen but

its occurrence is not constant and does not require taxonomie récognition.

Acknowledgements

I am grateful to Professor J. P. Troy for providing the collections which are the subject of

this paper and to Dr M. Bouché for transmitting them to me. Mr R. Raven is thanked for

measuring the setal ratios. Ail illustrations are by the author. This study was made possible

hv Australian Research Grants Committee support.

502

B. G. M. JAMIESON

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Manuscrit déposé le 7 septembre 1976.

Bull. Mus. natn. Hist. nat., Paris, 3 e sér., n° 450, mars-avril 1977,

Zoologie 313 : 477-502.

Achevé d’imprimer le 30 juillet 1977.

IMPRIMERIE NATIONALE

7 564 002 5

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Les références bibliographiques apparaîtront selon les modèles suivants :

Bauchot, M.-L., J. Daget, J.-C. Hureau et Th. Monod, 1970. — Le problème des

« auteurs secondaires » en taxionomie. Bull. Mus. Hist. nat., Paris, 2 e sér., 42 (2) : 301-304.

Tinbergen, N., 1952. — The study of instinct. Oxford, Clarendon Press, 228 p.

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