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Revision of the New Zealand Pill Millipedes (Oniscomorpha, Sphaerotheridae)

[Received by the Editor, October 5, 1955.]

Abstract

A revision of the N.Z Sphaerotheridae based on all the available specimens in N.Z. museum collections. Five forms are recognized: Procyliosoma (Procyliosoma) leiosomum (Hutton), Procyliosoma (Procyliosoma) tuberculatum tuberculatum Silvestri, Procyliosoma (Procyliosoma) delaeyi delacyi (White), Procyliosoma (Procyliosoma) delacyi striolatum (Pocock), and a new subspecies Procyliosoma (Procyliosoma) tuberculatum westlandicum. A key is given for separating the five forms, which are redescribed and figured. Detailed discussion of variation in P. tuberculatum tuberculatum and P. delacyi striolatum is given.

Introduction

The Sphaerotheridae are large-bodied millipedes which have thirteen body somites and twenty-one pairs of walking legs (Fig. 1). In the male, posterior to the last pair of legs, there are two pairs of telopods, each made up of a syncoxite and two telopodites which may be chelate. The apertures of the genital ducts are on the coxae of the second legs and in the female are surrounded by sclerotized laminae. The tergites are strongly curved and horny, and, except for the last, each is overlapped slightly by the preceding tergite. The first tergite, or collum, is very small; the last tergite, or pygidium, is large and semicircular in shape.

The Sphaerotheridae exhibit complete enrollment or conglobation, being capable of rolling the body into a ball so that the head and ventral structures are completely covered by the pygidium (fig. 2). The popular name of “pill millipedes” has arisen from the more or less spherical form assumed during conglobation and possibly also from the early practice in Europe of coating the enrolled millipede with sugar for sale as pills with alleged therapeutic properties. The enrollment habit is one of the most highly developed means of passive defence amongst the arthropods, the head and sense organs being completely protected, and the entire animal being extremely resistant to external pressure.

Pill millipedes have been collected from scattered localities throughout the North and South Islands of New Zealand, where there is an abundant supply of decaying leaves from endemic forests, especially beech. During the day they are found curled up under logs and stones, or buried in the ground, but at night they emerge to feed on decaying leaves.

The first pill millipede recorded from New Zealand was that described by White (1859) as Zephronia De Lacyi. Three other species, namely Sphaerotherium leiosomum Hutton (1877), S. novae-zealandiae Kirk (1885) and Cyliosoma striolatum Pocock (1895), were later described. The largest of the New Zealand species, Procyliosoma tuberculatum, was described in 1917 by Silvestri when he revised the Australian and New Zealand Sphaerotheridae. They were placed in two genera, Cyliosoma Pocock (1895) and a new genus Procyliosoma, which included all the New Zealand species and some of the Australian species. Procyliosoma was obviously erected without an examination being made of Sphaerotherium angulatum Butler, the type species of Cyliosoma, since Silvestri states that this and five other species.

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from the region require re-examination. The diagnostic characters which he used for separating Procyliosoma from Cyliosoma had not been given by Butler (1878) and Pocock (1895) in the description of S. angulatum.

Silvestri gave a fairly extensive diagnosis of Cyliosoma which he based on C. targionii Silvestri. He divided Cyliosoma into two subgenera, Cyliosoma s.s. and Epicyliosoma, again using characters which had not been considered in the original descriptions of the type species. In this division, C. angulatum (Butler) automatically became the type species of the subgenus Cyliosoma, although this is not stated in the paper; no type species was designated for Epicyliosoma.

Using the female genital plates and apical cones on the antennae as diagnostic characters, Silvestri divided Procyliosoma into two subgenera, Procyliosoma s.s. which included the New Zealand species and two new species from Australia, and Syncliosoma which was based on one female specimen from Australia.

In a key to the genera of the Sphaerotheridae, Attems (1928) has separated Procyliosoma Silvestri from Kylindotherium Attems, using mainly the form of the telopods. He states that in Procyliosoma the anterior telopod has no stridulating organ, and that the tibio-tarsus is undivided. For all the New Zealand species, and possibly the Australian species, this statement is incorrect. It probably arose from the inadequate diagnosis of Procyliosoma, and the poor figures and brief descriptions of the species given by Silvestri. If Attems' key were used, males of the New Zealand species of Procyliosoma would all be placed in Kylindotherium Attems, and so also would the females of at least one species.

To avoid further confusion in the taxonomy of this family, the classification of the Sphaerotheridae as given by Silvestri in his revision has been used in this paper However, it must be realized that the generic position of the Australian and New Zealand species cannot be definitely established until the type species of Cyliosoma has been carefully re-examined. It is quite possible that C. angulatum (Butler) possesses the characters of Silvestri's Procyliosoma, or, if it does belong to his Cyliosoma s.I., it may fall into his subgenus Epicyliosoma. Hence there seems to be no point at this stage in designating a type species for the latter subgenus.

This revision, which is the first undertaken by a New Zealand worker, deals with five forms of Procyliosoma. One of these is a new subspecies, and another, which previously ranked as a species, has been reduced to a subspecies. The remaining three species are redescribed and figured. The material examined comprises all the pill millipedes in the New Zealand museum collections, and in the Cawthron Institute collection.

Systematics

Family Sphaerotheridae Silvestii, 1896
Genus Procyliosoma Silvestri, 1917
Subgenus Procyliosoma Silvestri, 1917

Type Species, (o.d.) Procyliosoma (Procyliosoma) leae Silvestii, Hobart, Tasmania.

1917. Procyliosoma (s.s.) leae Silvestri, Boll. Lab. Zool. Portict, 12: 77–8, Fig. XI.

The New Zealand species of Procyliosoma form a compact group, having the following characters in common:

The antennae are 6-segmented, the 6th segment being flattened and disc-like and bearing usually 4 apical sensory papillae; the number of papillae varies slightly, being as high as 9 in some specimens of P. tuberculatum tuberculatum.

The surface of the tergites under high magnification is finely and shallowly reticulate, and within the cells of this network are numerous fine punctuations, approximately 0.01 mm in diameter. Attems (1928) regards similar punctures in Sphaerotherium as being probably the openings of hypodermal glands. In addition to these minute punctures, there are larger punctures, usually termed setiferous pits or punctures, since they may bear a seta. Setiferous punctures are large and saucer-shaped in P. tuberculatum westlandicum n.subsp and in P. delacyi delacyi, measuring up to 0.25 mm in diameter. In P. tuberculatum tuberculatum

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Figs 1–2—Procyliosoma (Procyliosoma) tuberculatum tuberculum Silvestri 1, Adult female 2, Conglobate adult female (Approx twice natural size)
Figs 3–13—Procyliosoma (Procyliosoma) letosomum (Hutton). 3, Left eye 4, Shield 5, Tergite sculpturing (anterior half) 6, Tergite sculpturing (posterior half) 7, Pygidium of male 8, 10th leg. 9, 2nd leg of female. 10, Anterior telopod (anterior aspect) 11, 2nd and 3rd segments of anterior telopodite (posterior aspect). 12, Posterior telopodite and syncoxite (anterior aspect). 13, Movable finger of posterior telopodite (posterior aspect). (All scales equal 0.5 mm except where otherwise indicated)

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and P. delacyi striolatum they are shallower and smaller, measuring up to 0.05 mm in diameter. Small punctures are also present in P. delacyi delacyi.

The second tergite, or shield, is the deepest tergite. It has a rounded rim which extends along the anterior, lateral and postero-lateral margins. This rim is separated from the rest of the tergite by a furrow which is narrow anteriorly, but widens laterally and posteriorly like the brim of a hat, and is therefore called the brim of the shield. The declivity connecting the brim with the main part of the shield bears laterally a number of small ridges or keels. The distal outer margin of the coxa of the walking leg is produced either into a flattened triangular or rounded lobe, or into a number of spiny processes. The lobes and spines are absent from the 2nd coxae of the female. The 6th segment bears a deeply pigmented spine, the upper spine, on the outer margin, at the base of the claw, and a row of similar spines, the lower spines, along the inner edge. The upper spine may be absent from the 2nd leg of the female Rarely it is absent or duplicated in the walking legs.

The telopods are composed of a syncoxite and a pair of 3-segmented telopodites. The anterior telopod is small, with the two halves of the syncoxite distinct. The anterior telopodite has the 1st segment short, broad, and nearly cylindrical. The 2nd segment is longer and narrower, with the distal internal angle produced into a long or short process, the fixed finger, which may bear tubercles. The 3rd segment is long and narrow, curving inwards towards the fixed finger, and bearing a row of ridged knobs along its distal posterior margin. The posterior telopod is much larger than the anterior one, and its telopodite is chelate. The syncoxite shows no sign of a median suture; distally it is produced into two small hairy horns, the tips of which open out into funnels. On its aboral surface the syncoxite bears a pair of hairy, cushion-like lobes, the bristle lobes. The 1st segment of the posterior telopodite is short, broad, and nearly cylindrical. The 2nd is very long with a large process, the fixed finger, arising from its distal internal angle, the fixed finger has a pale rounded protuberance at its base. The 3rd segment, the movable finger, is long, narrow, tapering distally, and curving towards the fixed finger, and with a few small spines and a small, pale, rounded protuberance midway along its concave internal surface. The posterior edge of the movable finger bears a row of ridged knobs. The furrowed knobs on the 3rd segments, and any tubercles on the fixed fingers of the telopods are considered to be stridulating organs in the Sphaerotheridae.

Silvestri distinguishes Procyliosoma from Cyliosoma by the form of the anterior telopodite. In Cyliosoma the fixed finger arises from the internal distal angle of the first segment, while in Procyliosoma it arises in a similar position from the second segment. Also, in Cyliosoma the claw of the walking leg is much more strongly curved. He also used the form of the laminae of the vulva as a generic difference, considering the external proximal lamina to be much larger than the internal one in Procyliosoma, whereas they are approximately the same size in Cyliosoma. Although this difference in the vulvae appears to be true for all the Australian species figured by Silvestri, the size of the laminae has been found to vary so much in one New Zealand species that the proximal laminae appear subequal. Silvestri's subgeneric division of Procyliosoma is based on the number of antennal sensory cones, and the form of the distal lamina of the vulva. Procyliosoma has 4 antennal cones, and the distal lamina is short with the inner margin rounded, while Syncyliosoma has numerous antennal cones and the distal lamina is rather long and subtriangular. In the New Zealand species there is slight variation in the number of antennal papillae, and there is considerable variation in the form of the distal lamina, so that it may be subtriangular.

Key to the New Zealand Species and Subspecies of Procyliosoma

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

1 Eye composed of 30–40 ocelli differing markedly in size, with 3 or 4 along the outer edge much larger than any others. Tergites uniformly black. 6th segment of leg robust, less than 4 × longer than wide
Eye composed of at least 50 ocelli, which are subequal in size. Tergites brown marbled with black. 6th segment of leg slender, at least 4 × longer than wide
2
4
2 Distal outer margin of coxa with 3 or more spiny processes
Distal outer margin of coxa without spines, instead produced into a flattened triangular lobe
P. leiosomum 3
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[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

3 Setiferous punctures small and shallow, not greater than 0.05 mm in diameter
Setiferous punctures large and saucer-shaped, about 0.2 mm in diameter
P. tuberculatum tuberculatum
P. tuberculatum westlandicum n subsp.
4. Tergites with scattered small setiferous punctures, and with large setiferous punctures on their anterior half
Tergites with dense small setiferous punctures only
P. delacyi delacyi
P. delacyi striolatum

Procyliosoma (Procyliosoma) leiosomum (Hutton, 1877) Figs. 3–13.

1877. Sphaerotherum leiosomum Hutton, Ann. Mag. Nat. Hist., ser. 4, XX: 116–7.

1878. Sphaerotherium leiosomus Hutton, Trans. N.Z. Inst., X: 291–2, Pl. XI, Fig. 6.

1895. Cyliosoma liosoma Pocock, Ann. Mag. Nat. Hist., ser. 6, XVI: 415.

1917. Procyliosoma (s.s.) leiosoma Silvestri, Boll. Lab. Zool. Portici, 12: 79–80, Fig. XIII.

Colour dark olive-black Head, collum, brim of shield olive-brown. Male width up to 7 mm, length up to 13 mm; female width up to 8 mm, length up to 15 mm.

Head with coarse setiferous pits, denser in the fore part. Eye as in Fig. 3, not very prominent, with approximately 33 ocelli, of which 2 or 3 along the outer edge are very large. Collum not pitted, but with a row of submarginal setae along the anterior and posterior margins Surface of tergites and shield finely reticulate anteriorly, changing to coarsely reticulate near the posterior margin, where there is a row of widely spaced longish setae. Anterior half to three-quarters of each tergite finely and densely punctulate, the punctures approximately 0.03 mm in diameter (Fig. 5). Shield as in Fig. 4, brim not broad; declivity with 8 to 10 weak keels; an irregular row of longish setae anteriorly and laterally in the furrow. Shield with a row of longish setae just inside the posterior margin, and a few longish setae scattered across the rest of the surface. Pygidium evenly rounded, descending rather steeply posteriorly; surface reticulate, very faintly pilose, with a few long submarginal hairs; margin slightly thickened; underside pubescent, with a moderately long, dark brown longitudinal keel on each side. The keel may be divided. Similar but much reduced keels on the underside of the 4th to 12th tergites. Pygidium of male as in Fig. 7, with a median elongate-oval whitish protuberance which has a few long setae on either side of it.

Leg as in Fig. 8, stout. Coxa with the distal outer angle produced into a flattened, approximately triangular, lobe which has a faintly serrated margin and bears as many as 6 marginal setae. 3rd segment stout. 6th segment short, thick, less than 4 times longer than wide, scarcely tapering distally so that the width halfway along its length is equal to that at the base; 6 to 9 lower spines; claw with the basal tooth rather sharp.

2nd leg of female as in Fig. 9.6th segment about 2 ½ tunes wider than long, with 6 or 7 lower spines but no upper spine. Vulva with distal lamina short, flattened and lacking setae. Outer proximal lamina larger than inner lamina, both with a few hairs distally.

Anterior telepods as in Figs. 10 and 11. Syncoxal segment with sparse setae. 1st segment of telopodite very hairy, especially along the inner edge; outer margin produced into a flattened triangular blade bearing a few marginal setae distally. 2nd segment narrower than 1st, with setae mainly along the outer margin and on the adoral surface. Fixed finger short, blade-like, tip serratulate; 1 to 3 spines on the adoral surface near the base of the movable finger. Movable finger much longer than fixed finger, tapering distally, hairy proximally and laterally, with 7 to 10 ridged teeth; 1 spine on adoral surface distally, and 1 to 3 spines on concave aboral surface The 2 fingers capable of forming pincers.

Posterior telopods as in Figs. 12 and 13, syncoxite rather broad, with scattered setae. Horns very short, rather wide; funnels wide. Bristle lobes subtriangular. Telopodite with 1st segment moderately hairy, 2nd segment very hairy, especially along outer edge. Fixed finger long and stout, lappet large; adoral surface with 1 spine and a median tract of 6 or 7 long setae. Movable finger long, hairy near base, with 9 to 12 ridged teeth and a very small papilla; aboral surface with 1 or 2 spines proximal to and 1 spine distal to the papilla.

Hutton described this species from material in the Otago Museum, giving the type locality as Dunedin. To date, the type specimen has not been located. Amongst the Otago Museum material examined by the writer is a series of specimens collected in Dunedin by G. M. Thomson, no date, and labelled Sphaerotherium leiosomus, possibly in Hutton's handwriting, but there is nothing to indicate that the type specimen is in this material. The matter is complicated also by the fact that although most of the specimens are Procyliosoma leiosomum, in the tube are 2 specimens of Procyliosoma tuberculatum tuberculatum stated to be from Dunedin, but which the writer considers to be probably from Cape Maria van Diemen. However, from.

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the descriptions and figures given by Hutton, the species can be fairly easily separated from the other New Zealand species. The specimens described and figured in this paper are from Notornis Valley, Te Anau. They are identical with the specimens of P. leiosomum from Dunedin, but are in a much better state of preservation.

Localities. Lake Te Au, south arm of Lake Te Anau, 12–24.1.1953, R. R. Forster, 1 ♀ (C.M. Myr. 46); Notornis Valley, middle arm of Lake Te Anau, 11.1.1949, J. H. Sorensen, 1 ♂ (D.M. 8/121); same locality, 20.2.1954, P. C. Bull, 5 ♂, 1 ♀, 1 imm. ♂ (D.M. 8/7–13); Otautau, Southland, 20.8.1954, O. Sanson, 2 ♀ (D.M. 8/14, 8/130); Ben Lomond, 2,000ft, Otago, 7.3.1943, J. T. Salmon, 1 ♀ (D M. 8/57); Dunedin, no date, G. M. Thomson, 1 ♂, 3 ♀, (O. M. Myr 11, 14–16); Tapanui, Otago, 25.1.1951, R. R. Forster, 3 ♀ (C.M. Myr. 26–28).

Procyliosoma (Procyliosoma) tuberculatum tuberculatum Silvestri, 1917. Figs. 1–2, 14–32.

1917. Procyliosoma (s.s.) tuberculatum Silvestri, Boll. Lab. Zool. Portici, 12: 80–1, Fig. XIV.

Colour blackish-brown or black. Head chestnut or dark brown, with the anterior margin black, and sometimes with a small median triangular ochreous area immediately behind the anterior margin. Male width up to 11 mm, length up to 22 mm; female width up to 20 mm, length up to 40 mm.

Head with coarse setiferous pits, moderately dense on the fore part, very sparse on the vertex. Eye as in Fig. 14, not very prominent, with approximately 40 ocelli of variable sizes, 3 or 4 along the outer margin being much larger than all the rest. Collum with a few long setae very widely spaced around the anterior and posterior margins. Surface of collum and other tergites finely reticulate, the network being coarser near the posterior border. Setiferous pits rather small, up to 0.05 mm in diameter (Fig. 16) dense or sparse all over tergite, or restricted to the anterior half of each tergite. Collum with these pits very sparse or absent. Tergites with a few marginal setae posteriorly, and in some specimens the 6th to 11th tergites with a lateral tubercle. Shield as in Fig. 15, brim not broad; declivity not steep, with 6 to 8 very feeble keels. In large specimens the keels may be absent. Setae scattered over furrow and declivity. Shield with a row of widely spaced hairs along the posterior margin, setae sparse or absent on the rest of the surface of the shield. Pygidium with the sides shallowly depressed; surface finely reticulate, faintly or densely pilose; margin slightly thickened, and projecting slightly in the middle, submarginal hairs sparse, underside very hairy with a dark-coloured longitudinal keel on each side anteriorly. Similar keels on the 4th to 12th tergites, those of the 9th to 11th being reduced to small knobs.

Leg as in Figs. 17 and 18, stout. Coxa with the distal outer angle bearing 1 or more (usually at least 3) simple spiny processes These processes may bear 3 or 4 short hairs which do not arise from near the tips. 3rd segment stout. 6th segment robust, about 3 ½ times longer than wide, scarcely tapering distally so that the width halfway along it is equal to that of the base; 5 to 9 lower spines; claw with the basal tooth rather wide and ledge-like.

2nd leg of the female as in Fig. 19. 6th segment approximately 2 ½ times longer than wide, with 4 to 6 lower spines, upper spine absent. Vulva with distal lamina flattened and bearing a few setae along the distal margin. Its shape varies from being approximately rectangular and about 2 ½ times wider than long, to being semi-circular or triangular Proximal external lamina considerably larger than internal lamina, both with scattered setae.

Anterior telopods as in Figs. 20 and 21, syncoxal segment almost naked. 1st segment of telopodite very sparsely setose, inner edge rather sharp and extending inwards well beyond base of 2nd segment. 2nd segment rather flattened, with a few setae distally Fixed finger very long, flattened, tip crenulate; adoral surface with 4 to 6 setae and 0 to 3 spines. Movable finger very long, tapering distally, almost naked, with 9 to 13 ridged teeth, 0 to 2 spines on the aboral surface. The 2 fingers form strong pincers.

Posterior telopods as in Figs. 22 and 23, syncoxite elongate and slender. Horns rather long, funnels elongate and narrow. Bristle lobes rounded, not very deep Telopodite very large. 1st and 2nd segment scarcely hairy. Fixed finger very long and thin, with 0 to 2 small spines and a tract of 6 to 10 long setae on the adoral surface; lappet large. Movable finger very long, slender, a few hairs near the base, 14 to 20 ridged teeth; a small papilla with 1 or 2 small spines proximal to it.

The density of setiferous pits on the surface of the tergites and pygidium shows considerable geographical. variation, but, in general, within one population it is fairly constant Variation in the actual size of the punctures is extremely slight throughout the entire sub-species. Specimens from Cape Maria van Diemen have the entire surface of the tergites and pygidium covered with fine close pits, with the setae minute Those from Tc Araroa and Whakarewarewa.

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Figs. 14–23—Procyliosoma (Procyliosoma) tuberculatum tuberculatum Silvestri. Specimens from Cape Maria van Diemen 14, Right eye 15, Shield. 16, Tergite sculpturing 17, 10th leg. 18, Coxal spines of 10th leg 19, 2nd leg of female 20, Anterior telopod (anterior aspect) 21, 2nd and 3rd segments of anterior telopodite (posteriot aspect) 22, Posterior telopodite and syncoxite (anterior aspect). 23, Movable finger of telopodite (posterior aspect). (All scales equal to 0.5 mm except where otherwise indicated).

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have similarly fine dense pits, but only on the anterior half to three-quarters of each tergite, and the pygidium is beset with dense, short, stiff hairs which are clearly visible at about 30 magnifications. In specimens from the Ruahine Range, the pits are fine but sparse, with minute setae, and are on only the anterior half of each tergite. The pygidium is not pilose. Setiferous pits are fine and dense on the anterior half to two-thirds of each tergite in specimens from the Tararua Ranges, Rimutaka Ranges, Orongorongo Valley, Kaitoke, Akatarawa, Stephen Island and Queen Charlotte Sound. In these specimens the setae are minute on the tergites and appear to be absent from the pygidium of all specimens except a few from the Rimutaka Range, in which they are very fine and sparse. In specimens from all localities within the Nelson area the pits are sparse. They may be slightly larger than those on specimens from Cape Maria van Diemen, and each pit may be supplied with a moderately long hair. The pits extend over the entire surface or are restricted to the anterior half to two-thirds of the tergite. With the exception of specimens from Punapaua, all specimens examined from the Nelson area have the main surface of the pygidium either naked, or with a few minute setae. In male specimens from Punapaua, the pygidium is densely covered with short, stiff hairs; in females the hairs are much less dense; both males and females have very sparse pits which are restricted to the anterior quarter of each tergite.

The shape of the distal lamina of the vulva is highly variable, sometimes even within individuals of one population. Some of the variations are shown in Figs. 24 to 32. The vulva is always oval, longer than wide, and not very densely pigmented. Characteristically, the distal lamina in this subspecies is flattened so that its main surface does not project beyond the level of that of the proximal laminae, and the distal margin tends to be sharp rather than rounded.

In general, there does not appear to be any correlation between sculpturing on the tergites and the shape of the distal lamina.

Silvestri described this species from specimens collected at Cape Maria van Diemen, Croixelles (Marlborough), French Pass, and Stephen Island, without designating a definite type locality. Specimens have been described and figured in this paper from Cape Maria van Diemen and this is here designated type locality.

Localities. North Island: Cape Maria van Diemen, no date, A. Hamilton, 3 ♂, 5 ♀ (O.M. Myr. 7–10, 17–20); Stephen Island, Cook Strait, no other data, 2 ♂, 2 ♀, 2 imm. ♀ (O.M. Myr. 1–6); Ruahine Mountains, 12.5.1939, A. C. O'Connor, 1 ♀ (D. M. 8/99); Tauherenikau Valley, Tararua Ranges, 8.2.1949, B. A. Holloway, 1 ♂, 1 ♀ (D.M. 8/133–134); slopes of Mt Tauherenikau, Tararua Ranges, 21.2.1952, B. A. Holloway, 1 ♂ (D. M.8/131); Pakuratahi Forks, Kaitoke, Wellington, 26.1.1954, W. J. Huggins, 4 ♀ (D.M. 8/92–95); Orongorongo Valley. 19.1.1953, S. Traill, 1 ♀ (D.M. 8/98); 5-mile Track, Orongorongo Area, 10.4.1954, L. Hampson-Tindale, 1 ♀ (D.M. 8/138); Rimutaka Range, western side, 800ft, 13.6.1948, R. K. Dell and A. C. O'Connor, 3 ♂, 5 ♀ (D.M. 8/73–79, 8/163); Whakarewarewa, Rotorua, 4.11 1954, R. Zondag, 10 ♂, 10 ♀ (D.M 8/167–169. 8/170–174, 8/184–195); same locality, 19.10 1952, G. B. Rawlings, 5 ♂, 3 ♀ (D M. 8/129, 8/135–136, 8/140, 8/159–162); Te Araroa, Waipu County, no other data, 3 ♀ (D.M. 8/4–6); Little Akatarawa, Wellington, 5.1.1922, F. W. Vosseler, 2 ♀ (C.I. Myr. 12, 13); Akatarawa, Wellington, 4.11.1952, M. Redington, 1 ♀ (D.M 8/158); same locality and collector, 13.4.1953, 1 ♀ (D.M. 8/139); same locality, M. Ordish, 10.10.1954, 1 ♀ (D.M. 8/128).

South Island: Dunedin, no date, G. M. Thomson, 2 ♂ (O.M. Myr. 12, 13); Punapaua, West Haven, Nelson, 17.3.1955, R. Ordish, 5 ♂, 1 ♀ (D.M 8/175–180); Beeby's Knob, 4,300ft; Nelson, 23.2.1954, E. S. Gourlay, 1 ♀ (D.M. 8/29); Nelson, 18.5 1924, E. S. Gourlay, 2 ♂, 3 ♀ (C. I. Myr. 2–6); same locality and collector,—.3.1954, 1 ♀ (C.I. Myr. 1); Pelorus Reserve, Nelson, 12.11.1949, A. W. Parrott, 1 ♀, 1 imm. ♀, (C I. Myr. 7–8); Canaan, Nelson,—.12 1950 and—.12.1951, A. C. O'Connor, 7 ♀ (D.M. 8/87–91, 8/96–97); Dun. Mt., Nelson, 8.3.1952, B. A. Holloway, 1 ♂ (D.M. 8/137); Maitai Valley, Nelson, 17.5.1952, B. A. Holloway, 1 ♂ (D M. 8/132); Gowan, Murchison, 2.1.1920, coll.?, 1 ♂ (D.M. 8/28); Queen Charlotte Sound, 10.3.1930, E. Walker, 1 imm. ♀ (C.M. Myr. 74); Resolution Cove, Queen Charlotte Sound. 12.1.1919, R. S. Allen, 1 ♀ (C.M Myr 6): Ship's Cove, Queen Charlotte Sound, 12.9.1948, R. K. Dell, 3 ♂, 2 ♀ (D.M. 8/82–86); Bluemine Island, Queen Charlotte Sound, 4.9.1948, R. K. Dell, 2 ♀ (D.M. 8/80–81).

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It is doubtful whether the subspecies still exists on Stephen Island. The specimens stated as being from Dunedin appear to be identical with the Cape Maria van Diemen specimens and probably are from that locality.

Procyliosoma (Procyliosoma) tuberculatum westlandicum n. subsp. Figs. 33–37.

Colour as in P. tuberculatum tuberculatum except that the yellowish triangular area on the head is more prominent. Female width up to 13 mm, length up to 28 mm; male width 9 mm, length 17 mm.

Head, collum and shield as in P. tuberculatum tuberculatum. The sculpturing of the tergites differs in that the pits are much larger and deeper, measuring up to 0.2 mm in diameter (Fig. 33). They are arranged in a wide, prominent band across the anterior half of the tergite, being largest and densest near the middle of each tergite. Their setae are fine. There are no tubercles on the sides of the tergites. Shape and underside of pygidium as in P. tuberculatum tuberculatum. Dorsal surface of pygidium covered with fine, dense, short hairs easily seen under 30 magnifications.

Leg as in P. tuberculatum tuberculatum except that the coxal spiny processes are very large, branched and usually bear long setae, some of which arise from near the tips of the branches (Fig. 35). 2nd leg of the female as in Fig. 36, 6th segment with 1 to 3 lower spines. Vulva very heavily pigmented; distal lamina short, very much thickened and with its distal margin rounded, not flattened.

Anterior telopods as in Fig. 37, differing from those of P. tuberculatum tuberculatum as follows 2nd segment of telopodite much more hairy, the hairs rather long Fixed finger shorter than in P. tuberculatum tuberculatum, with 2 long setae at the tip, and 1 spine and a few longish setae scattered over the adoral surface Movable finger much longer than fixed finger, its setae longer and more numerous; 4 to 6 ridged teeth.

Posterior telopods as in P. tuberculatum tuberculatum except that the base of the fixed finger is more hairy, and the tip crenulate. The hairs between the proximal edge and the papilla of the movable finger are much longer than those in P. tuberculatum tuberculatum There are 12 to 13 ridged teeth.

Types. Holotype female (Myr. 70 [tube], 70.1, 70.2 [slides]), allotype male (Myr. 1 [tube], 1.1, 1.2, [slides]) and 2 female paratypes (Myr. 5, 5.1, 71, 71.1, 71.2) all in the Canterbury Museum collection.

1 paratype female (8/203 and 9/218) in the Dominion Museum collection.

Localities. Buller Creek, Punakaiki, Westland, R. R. Forster. 24.1.1950 (type locality); Karamea Bluff, West Nelson, G. S. Southgate, 4.11.1950. (allotype): Seddonville, West Nelson, G. S. Southgate, 6.11.1950.

This subspecies appears to be confined to an area west of the Mt Arthur Tableland, but within the actual area the northern and southern limitations have not been determined It would appear that the Tableland has, until relatively recent times, acted as a barrier in separating this subspecies from tuberculatum, which, in the Nelson area, seems to be confined to the region to the east of the Tableland. A similar distribution on either side of the Tableland is found in 2 series of land snails of the genus Paryphanta, namely P. lignaria and its subspecies, and P. hochstetteri and its subspecies. The lignaria group, with 7 named subspecies, is distributed round Mokihinui and Seddonville, extending towards Karamea. West Nelson. Paryphanta lignaria oconnori Powell extends to the headwaters of the Leslie River, a tributary of the Karamea River, and to the western slopes of Mt. Arthur. On the eastern side of Mt. Arthur. Tableland, Paryphanta hochstetteri hochstetteri (Pfeiffer), a member of a polytypic form which is restricted to Northern and Eastern Nelson, extends as far east as Flora Camp (Powell, 1937, 1949, and Dell pers. comm.).

That the Tableland appears to be losing its effectiveness as a barrier for the 2 subspecies of tuberculatum is seen from the fact that hybrids have been collected from the Tableland area itself. The vulvae and the anterior part of the tergites of 2 of these hybrid specimens are shown in Figs. 38 to 41 Both specimens are from Salisbury's Clearing, Mt. Arthur. One specimen (Figs. 38 and 39) has punctures characteristic of both tuberculatum and westlandicum on the tergites and the vulva is flattened as in tuberculatum. The other specimen (Figs. 40 and 41) also has pits characteristic of both subspecies, but the smaller ones typical of tuberculatum predominate, and the larger ones are not quite as large as those of westlandicum. The.

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Figs. 24–32.—Procyliosoma (Procyliosoma) tuberculatum tuberculatum Silvestri. Variations in the shape of vulvae 24, Nelson specimen. 25–26, Canaan specimens. 27, Ruahine specimen. 28, Stephen Island specimen. 29, Whakarewarewa specimen. 30, Akatarawa specimen. 31, Punapaua specimen 32, Kaitoke specimen
Figs. 33–37—Procyliosoma (Procyliosoma) tuberculatum westlandicum n. subsp. 33, Tergite sculpturing (anterior half). 34, Tergite sculpturing (posterior half). 35, Coxal spines of 10th leg 36, 2nd leg of female 37, 2nd and 3rd segments of anterior telopod (anterior aspect)
Figs. 38–41.—Procyliosoma (Procyliosoma) tuberculatum × westlandicum. 38, Tergite sculpturing (anterior half) 39, Vulva of same specimen. 40, Tergite sculpturing (anterior half) 41, Vulva of same specimen (All scales equal to 0.5 mm except where otherwise indicated Figs. 24–32 drawn to same scale.)

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vulva is more characteristic of westlandicum than of tuberculatum. In both specimens the pygidium is very slightly pilose. The only male specimen in the series is from Salisbury's Clearing. It is very close to the nominate subspecies in the form of the telopods and tergites, but the pygidium is rather pilose. Only 1 hybrid is from the western side of the Tableland, namely from Leslie River Track. This specimen has the pygidium and vulva of tuberculatum but the tergite sculpturing is typical of westlandicum. The lower reaches of the Karamea River, and the Leslie River Valley, would possibly provide an easy migration route on to the Tableland for westlandicum.

Localities for P. tuberculatum tuberculatum × tuberculatum westlandicum. Salisbury's Opening, Mt. Arthur Tableland, 23.1.1948, R. R. Forster, 1 ♂, 1 ♀ (D. M.8/20, 8/21); Salisbury's Clearing, Mt, Arthur Tableland, 23.1.1948, R. K. Dell, 8 ♀ (D.M 8/112–117, 8/119, 8/120); Flora Track, Mt. Arthur Tableland, 7.4.1946, R. R. Forster, 1 ♀ (D.M. 8/71); Leslie River Track. Mt Arthur Tableland, 24.1.1948, R. K. Dell, 1 ♀ (D.M. 8/72).

Procyliosoma (Procyliosoma) delacyi delacyi (White, 1859), Figs. 42–50.

1859. Zephronia (Sphaerotherium) De Lacyi White, Ann. Mag. Nat. Hist., ser. 3, III: 406, Pl VIII, Figs. 2, 2a.

1873. Sphaerotherium de lacyi Butler, Proc. Zool. Soc. London: 177.

1895. Cyliosoma de lacyi Pocock, Ann. Mag. Nat. Hist., ser. 6, XVI. 415.

1885. Sphaerotherium novae-zealandiae Kirk, Trans. N. Z. Inst., XVIII. 139–40, Figs. 1–3.

Colour (in alcohol) light brown-olivaceous In some specimens all or some of the tergites marbled or spotted with black Male width up to 8 mm. length up to 15 mm female width up to 9 mm, length up to 17 mm.

Head with coarse setiferous pits moderately dense in front, sparse on the vertex Eye as in Fig. 42. very large, compact, raised well above the surface of the head, composed of about 60 subequal ocelli arranged more-or-less in rows. Collum finely reticulate with dense small pits, a few widely spaced long setae just inside the posterior margin Surface of 3rd to 11th tergites as in Fig. 44, finely reticulate, with scattered small pits up to 0.03 mm in diameter and much larger saucer-shaped pits (up to 0.25 mm in diameter) mainly on the anterior half Shield as in Fig. 43, finely and densely punctulate, with a few large setiferous pits scattered over the surface but not along the posterior margin. Brim fairly wide; declivity steep, with 7 to 9 narrow keels which arise from a rounded rim at the top of the declivity; furrow deep with a row of setae extending into the brim. Pygidium wide, much shorter dorso-ventrally than the other tergites, not descending steeply; margin thickened and raised like the edge of a bell. On each side a shallow emargination separates off from the rest of the pygidium an anterior section about the width of the 12th tergite. Posterior margin of the male with or without a shallow median emargination. Underside of pygidium pilose, and on each side a small brown knob, with a very short, wide longitudinal keel immediately posterior to it. A much reduced keel on the underside of the 4th to 12th tergites.

Leg as in Fig. 45, long and slender. Coxa with the distal outer angle produced into a flattened semicircular lobe which has its margin serrate; lobe with 1–7 submarginal setae. 3rd segment long and slender. Distal inner margin of 5th segment with or without a small brown spine. 6th segment long, slender, tapering distally, usually more than 4 times longer than wide; 8 to 11 lower spines; upper spine present or absent; claw with a rounded tooth at the base.

2nd leg of the female as in Fig. 46 6th segment about 4 times longer than wide; upper and lower spines as in other legs Vulva elongate-oval, not very hairy: distal lamina triangular, surface depressed proximally; proximal lamina not greatly larger than internal lamina.

Anterior telopods as in Figs. 47 and 48. Syncoxal segment with a few setae. 1st segment of telopodite wider than 2nd, rather hairy, distal half of outer edge blade-like. 2nd segment wide at the base, outer proximal edge blade-like: 1 to 3 very long setae arising near the proximal internal margin on the aboral surface Fixed finger minute, wedge-shaped, with 0 to 3 spines near the base, 0 to 3 hairs; tip faintly crenulate Movable finger short, broad, with very few hairs; distal margin truncate with 5 to 8 ridged teeth. Concave surface with 2 spines. The movable and fixed fingers do not form pincers.

Posterior telopods as in Figs. 49 and 50. Syncoxite robust, hairy distally. Horns and funnels very short and wide Bristle lobes small and rounded Telopodite massive. 1st segment with a few hairs. 2nd segment covered with short, fine hairs, hairs rather sparse on outer margin; 2 or 3 long setae on the aboral surface near the base of the fixed finger. Fixed finger short, thick, with a tract of fine, short hairs and 0 to 2 spines on the adoral surface; papilla large.

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Movable finger long, finely hairy near the base; papilla large with 1 or 2 spines at its base on the adoral surface; 8 to 13 ridged teeth.

Localities. Nihotapu, Auckland,—.3.1923, A.V.M., 2 ♂, 1 ♀ (D.M. 8/58–60); Watershed Reserve, Auckland, 26/2/1944, G. Chamberlain, 3 ♀ (D.M 8/50–52); Mangapehi, Auckland,—.11.1930, coll.?, 1 ♀ (A.M. Myr. 1); Patea, Taranaki, 8.8.1954, G. W. Ramsay, 1 ♀ (D.M. 8/15).

E. Browning, of the British Museum (Natural History) has kindly examined the type specimen (female) of Zephronia delacyi White and verified White's description of the tergite puncturing. Unfortunately, no type locality other than “N.Z” is given.

Procyliosoma (Procyliosoma) delacyi striolatum (Pocock, 1895), Figs. 51–59.

1895. Cyliosoma striolatum Pocock, Ann. Mag. Nat. Hist., ser. 6, XVI: 414–5.

1917. Procyliosoma (s.s.) Delacyi Silvestri (non White 1859), Boll. Lab. Zool. Portici, 12: 82–3, Fig. XV.

1917. Procyliosoma (s s) striolatum Silvestri, Boll. Lab Zool. Portici, 12: 83–4, Fig. XVI.

Colour as in P. delacyi delacyi. Marbled specimens frequently with the pygidium brownishochreous with or without black markings Non-marbled specimens usually with the hind margin of the tergites blackish. Male width up to 9 mm, length up to 18 mm, female width up to 10 mm, length up to 19 mm.

This subspecies differs from P. delacyi delacyi in the following characters: The collum, shield, tergites and pygidium are finely and densely punctulate, the punctures measuring up to 0.03 mm in diameter (Fig. 51). There are no larger punctures. The tergites have a few widely spaced longish setae a short distance in from the posterior margin. The pygidium is faintly pilose, with a few large submarginal setae.

Legs as in delacyi delacyi. The distal prolateral lobe of the coxa with or without marginal setae. In general, there appears to be some correlation between the colour pattern on the terg♂tes, and the number of marginal hairs on the coxal lobe. The lobes are usually very hairy (Fig. 52) in specimens which have the tergites distinctly marbled, whereas in almost all specimens having tergites without marbling, or with only a few spots, the coxae are devoid of marginal hairs. Specimens with or without marbling on the tergites occur within the same locality, but in the study material available from certain localities there appears to be a marked preponderance of specimens having a uniform colour pattern For instance, of 43 specimens from the Caswell Sound and Te Anau areas, 41 have marbled tergites and all specimens have from 1 to 16 hairs on at least one coxal lobe of the 10th legs There are only 2 specimens without hairs on one coxal lobe, and these specimens are distinctly marbled. All except 2 of the 23 specimens from Mt. Arthur Tableland, Nelson, have no marbling on the tergites, although some have a few small black spots, and there are no marginal hairs on the 10th coxal lobes The 2 marbled specimens have 1 to 3 marginal hairs on the corresponding lobes Of 47 specimens from Westland, the tergites of 24 are distinctly marbled and the coxal lobes of the 10th legs have from 0 to 12 marginal hairs, 6 specimens having either one or both coxae without marginal hairs. The tergites of the remaining 23 specimens are either not marbled, or are only finely spotted, and the coxal lobes of the 10th legs in these specimens have from 0 to 11 submarginal setae. Fifteen of these specimens have one or both lobes devoid of marginal hairs Figure 59 shows the apparent correlation between colour pattern and setae on the coxal lobe In all other respects the two colour forms appear to be similar.

The shape of the laminae of the vulvae, especially the distal laminae, is extremely variable even within the specimens from one population Some of the variations are shown in Figs. 53 to 58. In some specimens (Figs. 53, 54, 57. 58) the entire vulva appears to have been twisted laterally so that the aperture between the proximal laminae extends diagonally across the coxa instead of being more or less vertical. The form of the vulva does not appear to be related to colour form.

Anterior telopods as in delacyi delacyi except that some specimens have a small spine on the inner distal angle of the 1st segment of one or both telopodites. The movable finger has as many as 9 ridged teeth.

Posterior telopods as in delacyi delacyi except that the movable finger may have as many as 3 spines near the papilla, and as many as 14 ridged teeth.

It is possible that similar variation in the number of spines and teeth of the telopods will be found to exist in delacyi delacyi when further specimens are examined.

Localities. Nelson,—.1.1925, E. Richardson, 1 ♀ (D.M 8/27); Woodpecker Bay. Nelson, 22.1.1950, R. R. Forster, 1 ♀ (C. M. Myr. 7); Onakoka, Nelson, 21.10.1938, coll.?, 1 ♀ (D.M. 8/61); Oparara, Nelson, 26.1.1950, R. R. Forster, 3 ♀ (C.M. Myr. 2–4): Maruia Saddle. Nelson, 18.11.1952. G. W. Ramsay, 1 ♂,

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Figs. 42–50—Procyliosoma (Procyliosoma) delacyi delacyi (White) 42, Left eye 43, Shield 44, Tergite sculpturing 45, 10th leg 46, 2nd leg of female 47, Anterior telopod (anterior aspect) 48, 2nd and 3rd segments of anterior telopodite (posterior aspect) 49, Posterior telopodite and syncoxite 50, Movable finger of posterior telopodite (posterior aspect)
Figs 51–58—Procyliosoma (Procyliosoma) delacyi striolatum (Pocock) 51, Tergite sculpturing. 52, 10th leg 53–58, Variations in the shape of vulvae 53, Moana specimen 54–55, Mt Davie specimens 56, Mt Arthur specimen. 57–58, Lake Te Au specimens (All scales equal to 0.5 mm except where otherwise indicated Figs. 53–58 drawn to same scale.)

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2 imm. (D.M. 8/156–157); Canaan Track, Nelson, 17/1.1/1 948, A Carson, 6 ♀ (C.I. Myr. 9–11, 14–16); Dun Mt., Nelson, 7.3.1952, B. A. Holloway, 2 ♂, 1 ♀ (D.M. 8/164–166); Mt. Arthur, Nelson, 22.1.1948, R. K. Dell, 3 ♀ (D.M 8/45–47); same locality, 28.1.1948, R. R. Forster, 1 ♀ (D.M. 8/19); Salisbury's Clearing, Mt. Arthur Tableland, 23.1.1948, R. K. Dell, 1 ♀ (D.M. 8/118); Leslie River Track, Mt. Arthur Tableland, 24.1.1948, R. K. Dell, 2 ♂, 4 ♀ (D.M. 8/39–44); Mt. Arthur Track, R. K. Dell, 22.1.1948, 2 ♀ (D.M. 8/16, 8/17); same locality and date, R. R. Forster, 1 ♀ (D.M. 8/25); same locality and collector, 28.1.1948, 7 ♀, 4 imm. (D.M. 8/18, 8/196–202); Flora Track, Mt. Arthur, 7.4.1946, same collector, 7 ♀ (D.M. 8/64–70); Mt Davie, near Greymouth, Westland, 18.1.1950, R. R. Forster, 4 ♀ (C.M. Myr 8–11); Balloon Hut Bush, Mt Arthur Tableland, 20.2.1946, J. T. Salmon, 1 ♂ (D.M. 8/49); Rugged Burn Valley, West Te Anau, 15–18.2.1953, G. W. Ramsay, 1 ♂, 2 ♀ (D.M. 8/1–3); Lake Peel, Mt. Arthur Tableland, 19.2.1946, J. T. Salmon, 1 ♀ (D.M. 8/24); Lower Hollyford Valley, Otago, 27.12.1944, J. T. Salmon, 1 ♂ (D.M. 8/26); same locality, 2.2.1946, R. R. Forster, 1 ♂ (D.M. 8/48); Key Summit, Otago, 25.1.1946, R. R. Forster, 1 ♂ (D. M.8/53); Lugarburn, Te Anau, 20.5.1955, K. H. Miers, 3 ♀ (D.M. 8/181–183), Te Anau, 15.2.1953, G. W. Ramsay, 6 ♀ (D.M. 8/141–146); Cascade Creek, Te Anau, 25.1.1951, R. R. Forster, 1 ♀ (C.M. Myr 23); Lake Te Au, south arm of Te Anau, 12–24.1.1953, R. R. Forster, 2 ♂, 10 ♀ (C.M. Myr. 44–45, 47–56); Stillwater Base Camp, Caswell Sound, 13.3.1949, R. K. Dell, 3 ♂, 6 ♀ (D.M. 8/147–155); Upper Stillwater Camp, Caswell Sound, 11–12.4.1949, R. R. Forster, 1 ♂, 4 ♀ (C.M. Myr. 15, 17–20); Caswell Sound, 18.4.1949, R. R. Forster, 1 ♂, 1 ♀ (C. M. Myr. 16, 24); Leslie Clearing, Caswell Sound, 15.3.1949, R. K. Dell, 2 ♂ (D.M. 8/22–23); Waiho Gorge, Westland, 20.2.1927, A. Castle, 5 ♂, 1 ♀ (D.M 8/33–38); south bank of Poringa River, South Westland, 3.6.1954, J. M. Moreland, 1 ♂, 2 ♀, 1 imm. ♂, 2 imm. ♀ (D.M. 8/122–127); Lake Kanieri, Westland, 1.1.1950, J. H. Sorensen, 1 imm. ♀ (D.M. 8/62); Hari Hari, Westland,—.12.1920, C. E. Foweraker, 1 ♀ (A.M Myr. 2); Marsden Rd., Greymouth, 5.3.1949, P. Hughson, 3 ♀ (C.M. Myr. 12–14); Bullock Creek, Punakaikai, Westland, 24.1.1950, R. R. Forster, 1 ♀ (C.M. Myr. 73); Old Taylorville Rd., Westland, 6.11.1950, R. R. Forster, 1 ♂ (C.M. Myr. 25); Hector, Westland, 28.2.1948, A. Goddard, 1 ♂ (A.M. Myr. 17); between Aickens and Otira. 8.1.1949, E. W. Dawson, 2 ♀ (C.M. Myr. 21, 22); Chapman's Reserve, near Bold Head, Westland,—.12.1921, C. E. Foweraker, 1 ♂ (A M. Myr. 3); Otira, Westland,—3.1945, T. P. Harris, 3 ♀ (D. M.18/30–32); same locality, 13.1.1951, E. W. Dawson, 2 ♀ (C.M. Myr 29, 30); Moana, Westland, 10.3.1950, R. R. Forster, 1 ♂, 5 ♀, 1 imm ♂, 6 imm. ♀ (C. M. Myr. 31–39); Okarito, Westland, 7.12 1949, R. R. Forster, 3 ♂, 6 ♀, 2 imm ♂, 2 imm. ♀ (C.M. Myr. 57–69).

E. Browning, of the British Museum (Natural History) has advised the writer that a female specimen having the characteristics of the specimens described above as P. delacyi striolatum bears two labels, Cyliosoma sharpi Pocock and ? Cyliosoma striolatum Pocock, the latter label not being in Pocock's writing, and has the register number 1889.11.6.1 This specimen is not designated as a type but it appears to be the only labelled specimen of P. delacyi striolatum in the British Museum collection.

Discussion

The New Zealand Sphaerotheridae belong to two groups, namely the tuberculatum leiosomum group and the delacyi group. The former group includes P. leiosomum and P. tuberculatum tuberculatum and P. tuberculatum westlandicum, all of which have black tergites, the eye with a small number of ocelli of various sizes, and the 6th segment of the leg stout. The delacyi group includes P. delacyi delacyi and P. delacyi striolatum, two very close subspecies having brownish tergites which may.

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Fig 59—Histograms showing possible relationship between marbling of the tergites and marginal setae on coxal lobe of the 10th left leg in specimens of Procyliosoma (Procyliosoma) delacyi striolatum. The relationship is shown more clearly in the percentage histogram (inset).

be marked with black, the eye with a large number of subequal ocelli, and the 6th segment of the leg slender.

P. tuberculatum tuberculatum is distributed throughout the North Island, on some islands in Cook Strait, and in Nelson to the east of the Mt. Arthur Tableland. In West Nelson and Westland it has developed as a distinct subspecies, P. tuberculatum westlandicum. There is some intermingling of these two subspecies in the Mt. Arthur Tableland area. In Southland and Otago, P. tuberculatum tuberculatum is replaced by the much smaller P. leiosomum.

Locality records for P. delacyi in the North Island are too few to allow the southern limitations of delacyi delacyi and the northern limitations of delacyi striolatum to be determined. To date, delacyi striolatum is known definitely only from the South Island and delacyi delacyi only from the northern half of the North Island Within each of these two subspecies there are two colour forms, the colour pattern being, in general, correlated with the number of marginal hairs on the lobes (Fig. 59).

From the very considerable variation shown in the puncturing of the tergites and form of the vulvae in P. tuberculatum tuberculatum, and in the tergite colouring, and form of the vulvae in P. delacyi delacyi, it would appear that these two forms are still in the very early stages of speciation.

Acknowledgments

This work was carried out at the Dominion Museum, Wellington, and the writer is grateful to R. K. Dell, of the Museum, for valuable discussions and criticism. Thanks are expressed to the directors of the Auckland, Canterbury, and Otago Museums, and of the Cawthron Institute, for the loan of specimens. The writer is also indebted to V. J. Wilson for assistance in the preparation of the paper and for seeing it through the press.

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References

Attems, C., 1928. The Myriopoda of South Africa. Ann. S. Afr. Mus., XXVI: 1–431.

Butler, A. G., 1878. Descriptions of Several New Species of Myriopoda of the Genera Sphaerotherium and Zephronia. Trans. Ent. Soc. London : 299–302.

Powell, A. W. B., 1937. “The Shellfish of New Zealand.” Auckland, Unity Press. 100 pp.

Powell, A. W. B., 1949. The Paryphantidae of New Zealand Rec. Auck. Inst. Mus., 3 (6): 347–67.

Miss B. A. Holloway

,
Biological Laboratories
, Harvard University,
Cambridge 38, Mass., U.S.A. (On leave from the Dominion Museum. Wellington.)