ON THE CLASSIFICATION OF THE FAMILY MACROCHELIDAE WITH PARTICULAR REFERENCE TO THE SUBFAMILY...

345

ON THE CLASSIFICATION OF THE FAMILY MACROCHELIDAE WITH PARTICULAR REFERENCE TO THE SUBFAMILY

PARHOLASPINAE (ACARINA-MESOSTIGMATA)

BY

G. OWEN EVANS Departvree,it of Zoology, Britask ilfuseicm (Natwal History)

[Accepted 13th March l95ti]

(With 64 figures in the text)

CONTENTS

. .

. .

. .

. . . . . . .. . . . . . . . . .. . . . . . . .. . . . . . . .. . . . . . . . . .. .. . . . .

page . . 346 . . 346 . . 360 .. 352 .. 353 .. 354 .. 357 . . 367 . . 359 . . 35!J . . 361 . . 36.3 .. 364 .. 385 . . 365 .. 366 . . 366: . . 366 . . 367 . . 369 . . 370 . . 371 . . 372 . . 373 .. 374 . . 376 . . 376 . . 377

IKTRODUC'TION

No comprehensive revisionary study of the classificatiori of the Macrochelidae has been published since Berlese's important work in 1918 and the present position is chaotic. This is largely due to the fact that, the original descriptions of many species, some of them genotypes, are so inadequate t'hat they cannot be

346 G . OWEK EVANS

recognized with certainty. Turk (1948) stressed trhis in his review of tjhe family in which he arranged the genera in four groups, mainly on the basis of the chaetotaxy of the ventri-anal shield in females.

The only other major revisionary works are those of Tragardh (1949 & 1952) in which proposals were made for the division of t,he family into subfamilies, but which, unfortunately, ignored Berlese’s fundamental work. Partial revisions are t8hose of Valle (1953) in his consideration of the genus GehoZaspis Berl. and Evans & Browning (1956) who have revised the British species of the subfamily Macrochelinae.

The present review of the classifica,tion of the Macrochelidae is based partly on an examination of material in t,he collections of the Brit’ish Museum (Nat. Hist.) and partly on the re-examination of some of the types of Berlese’s genera in the Stazione di Entomologia Agraria, Florence.

The nomenclature for the external morphology of the group follows, in the main, that used by Evans (1956).

.

CLASSIFICATION

The earliest attempt at a subfamily classification of the Macrochelidae was that proposed by Tragardh (1949) in which the family was divided, as follows, in two subfamilies :

” 1. Epistoinr a triaiigular iiiuoro without appendages. Legs I with ambulacres, metasteriial shields connected wit>h the sternal shield t,hrough a narrow bridge . . . . . . . . . . . . . . . . . .

Legs I without aiubulacres. Metasteriial shields distinct,, n u t fused wit’h the sternal shield . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . subfam. Macrochelilme.”

subfain. Proloholaspiiiae. -. Epistome with appendages.

In 1952, Tragardh modified his earlier classification and proposed a third subfamily, Areolaspinae, for Areolaspis Tragardh 1952 :

“ 1 . Metasteriial shields connected wit,h t>he sternal shield through a narrow bridge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Protolidaspinat Traghrdh, 1949.

-. Metasternal shields free . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. 2. Peritrematic shields not fused with the exopodal shields. Macrochelinae TrBgBrdh., 1949, -. Peritrematic shield8 fused with t,he exopodal shields . . . . Areolaspinae subfam. nov.’’

Evans & Browning (195ti) considered that, on the basis of the structure of leg I and the chelicerae, Protoholaspis Tragardh was not a macroclielid. Thus, the family was reduced to the t.wo subfamilies separated in couplet 2 of the above key. If Tragardh’s criterion for separating the Macrochelinae and Areo- laspinae, namely, the degree of fusion of the peritrematal and exopodal shields, is accepted then other genera, such as Gamasholaspis Berl., Holaspulus Berl., Purholaspis Berl., Euholcelaeno Bed. and Calholaspis Berl., must also be included in the Areolaspinae since all have the peritrematal shield partially or completely fused with the exopodal shield. An examination of the above genera as well as a number of new genera has revealed that the degree of fusion of the shields covering the venter of the female is considerably more varied than

supposed by Trligkrdh. below :

The possible fusions wit'hin this group are set out

I. perit reinstal-exopodal shields tiee. a. Metasternal shields and genital shield free (Fig. I a).

Purholuspis, Edholocelaeno, Gair~asltolu~pis, A rsoluspiss am1 Porlioluspiuu4ltrs gen. n.

b. Metasternal shields fused with the sterile1 ; genital shield free (Fig. 1 b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cafholnspis.

Peiitrematal-exopodal shields fused with the ventri-anal. t ~ . Metasternal and genital shields free (Fig. 1 c) . . . . . . . . . . Tricholaspis gen. n.

11.

Vig. 1 a-e.-Sclerotization of the venter of females of the snbfmnily Parholaspinae. u, Parholuspi~ Berl. : h, Cn!Iiolaspis Bed. : c, 7'richoln.spi.s~ g m . 1 1 . ; (1, N p . p ~ ~ , t h h p i k g ~ n . n. ; e, Holu-?puhL39 Berl. end. sh., endopodal shield : gem. ski., genital shield : iiwt. sh., inc+astenial shield ; per., peritreme ; per. sh., perit,reinatai shit=td ; st.., stigina. ; st. yh., st,eni;tl shield.

348 C. OWEN EVANS

b. Metasternal shields fused with the sternal ; genital shield free (Fig. 1 d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

a. Met&ernal shields free; genital shield fused with the ventri-anal (Fig. 1 e) ................................

Nep~irhoLu.s-pis geu. n.

Holasplus .

It is apparent, from the above, that the fusion of the ventral shields within t,he " Areolaspinae " can only be considered of secondary importance in the phylogeny of the family ; the importance given to this character in previous classifications being due to the fact that they provide simple key characters. On the other hand, the writer feels that such characters as the form of the peritreme, structure of the geiutal region, and to a lesser extent, the chaetotaxy of the palptarsus and the form of the tectum, are of fundamental importance in developing a natural classification of the family.

lat.sc1,

4

2 Figs. 2 4 - 2 . Looped peritreme of the Mecroclielinae. 3. Straight peritreme of the Parholaspinee.

4. Genital shield and lateral sclerites in Mucroclieles cnriciutu~ (C. L. Koch). 5. Palptarsus

gen. ah., genital shield ; lat. scl., lateral sclerite ; per., peritreme ; per. sh., peritremattll shield ; at. stigma.

O f a MWOChekS Sp.

Within the Macrochelidae there are two basic types of peritreme : one type . in which it is developed into a U-shaped loop and leaves the stigma posteriorly

CLASSIFIC.4TfON OF THE MAC’ROC‘HELIDAE 349

(Fig. 2) and the other, it1 wllich it issues from the anterior margill of the stigma without trace of a loop (Fig. 3). Other characters which are correlated with these different types of peritreme are :

I . The peritrematal shield is free in its posterior two-thirds when the peritreme is looped. When, however, it is straight, the peritrematal shield is fused with exopodal shields. The only exceptions are seen in Areo- laspis and Euholocelaeno where although the peritreme is looped the peritrematal shield is fused with the exopodal.

2 . In those genera with a looped peritreme, the genital shield is provided . with lateral sclerites (Fig. 4). The genital shield functions inde-

pendently of the sclerites and one is tempted to homologize them with the parasternal shields of the Parasitidae. Lateral sclerites are absent in the species with a straight peritreme.

3. The palptarsus is provided with a long rod-like seta (Fig. 5 ) externally in its distal third in a number of genera with a looped peritreme e.g. Macrocheles, Areoluspis and Holostmella. This characteristic seta has not been observed in the group with a straight peritreme.

4. The tectum is produced medially into three processes, which may be free or partly fused with each other, in those genera with a looped peritreme. When the tectum is straight, its anterior margin is denticulate and is with or without a simple median process. The only exceptions are Geholuspis Berl. s. lat. and Neopodocinurn Oudemans in which the tectum has a single median process but a looped peritreme, and Purholas- pulw in which there are lateral processes in addition to a single median process although the peritreme is straight.

The above group of correlated characters permits of the subdivisioii of the family into two groups of genera which are not associated with different environmental conditions. The characters which the members of each group have in common must be presumed, therefore, to be their common inheritance rather than common adaptive modifications. The two groups accordingly represent two phylogenetic assemblages within the family and are given subfamily status.

Family MACROCHELIDAE Vitzthum

Macroohelidae Vitzthum, H. G. (1930) Zool. Jnhrb. (Syst.) 59, 300.

Dorsal shield entire, variously ornamented, and usually with twenty-eight to thirty-two pairs of setae. Females with the usual ventral shields which show varying degrees of fusion wit,h each other ; genital shield with or without lateral sclerites. Male with separate ventri-anal or anal shield, or a holoventral shield ; genital orifice praesternal. Gnathosoma normal, salivary styli and corniculi well developed ; tectum simple or produced into a single or trifid process ; pedipalp with five free segments, specialised seta on the inner basal angle of the tarsus with three prongs. Chelicerae chelate-dentate and with a

350 Q . OWEN EVSNB

stout ventral seta in addition to the extero-ventral brush of setae ; movable digit with the spermatophoral process free distally. Legs I in both sexes without ambulacra : legs 11-IV with two claws and a lobate pulvillus. Legs I1 and sometrimes legs SV in the male, and rarely legs I1 in the female. with spurs. Free-living in decaying vegetation or paraphages of insects.

Key to subfamilies

I . Peritreme with a U-shaped loop atid leaving stigma posteriorly ; genital shield with lateral sclerites. . . . . . . . . . . .

- . Peritreme straight arid leaving stigma anteriorly ; genital shield without lateral sclerites. . . . . . . . . . . . . . . . . . . . . . . . . .

Macrocheliiiau Trgdli.

Parholaspime subfam. nov.

Subfamily MdCKOCHELINAE Tragirdh

This subfamily contains the majority of the described species 01 the There appears to have been a t least t,hree lines of evolution as indicated

The free-living Geholaspis group occupies an isolated position relative

Tricholocelaeno

family. in Fig. 6.

Euholocelaeno Holaspu,us

Fig. 6,-l)iagrainmat,ic representation of the pliyloge11y of thc Macrochelidao.

to the other Macrochelinae, being characterized by the form of the tectum and corniculi, and the tendency for increased sclerotizatioii of the ventri-anal region. The Areolaspis group although closely related to the Macrocheles group, is characterized by t'he development of the anterior region of the dorsal shield into a prow-like structure and by the armature of legs I1 in the female. The Macrocheles group is the most diverse ecologically and morphologically.

CLASSIFICATION OF THE MACROCHEWDAE 35 1

The genus Macrocheles contains a number of noii-specialized free-iiviiig species as well as paraphages of insects. The main trend of evolution in this group seems to be towards the development of specialized forms associated with Coleoptera. The structural modifications associated with the change from a free-living to a " parasitic " mode of life conceals the true relationship between the two groups and adds to the difficulty of developing a natural claesification. The genus Neopodocinum is an example of this.

The generic classification of the Macrochelinae is almost entirely based on the work of Berlese (1904 & 191%). Many of the genera have been in- sufficiently defined to be certain of their systematic position. In the following attempt a t a key to the genera, Trichocelaeno Berl. has been omitted since it is known from the male only. The lack of knowledge of the structure of the male throughout the subfamily results in the classification being based on the female.

Key to the genera of the Macrochelinae (Females only)

1. Vertical srtae arising from aii aiiterior I)i-oject,ioii of t,Iiu dorsal shield (Fig. 7 ; femur and often other segineiitv uf leg I1 with derotieed ridges or spurs .................... 3.

- . Vertical setae iiot arising from an anterior projection of the dorsal shield (Fig. 8); femur of leg I1 wit,hout ridges 01% spurs (except AndrhoIa.+.s). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Peritrernatrtl shield fused with the exopodal shield. 2.

EwholocPIae?zo Berl., 19 18 & -4wolospis TriigBrdh, 1952.

4. With anal shield. . . . . . . . . . . . . . . .

branched (Fig. 9 b) apically ; corniculi long and sleiider ; ventri-anal shield with five pairs of preanal setae. . . . . . . . . . Ge/w/ospis Berl., 1918 s. lat.

- . Tectuni with three anteriorly directed processes of about equal leiigth, which may he free (Fig. 10 a) or part>ly fused with each other (Fig. I 0 b) ; corniculi of normal lriigt,h ; ventri-anal shield normally with t,wo or three pairs of preaml

deeply oinarginate. In nests of bees of the geiius T r i p n i u . . . . . 1 rtyo?dio/trspis Vitzthuiu, 1930. -. Anterior iiiargiii of the steriial shield truricate or slightly

. . . . . . . . . . . . . . . . . . 8.

r l .

7. Ventri-anal

(including Holoce1uen.o Berl., 1910). - . Legs I & I1 with sclerotized ridges or spurs . . . . . Andrliokispis Turk, 1948.

* Provisionally, t,he writer considers the followiiigtJo be synonyms of rllncroclreles ; C'oprholnsppis Berl., 1918; Nothrolospis Berl., 1918; Dissoloncha Falconer, 1923; Mo?i.oplites Hull, 1925 and Neholaspis Turk, 1948.

352 G . OWEN EVANS

A 8

10 9 Figs. 7-10.-7. Anterior region of the dorsal shield of dreolaspi.9 b$~liolintus TriigBrdh. 8. Anterior

9, Tecta of Geholaspis s. str. (a) and Longi- 10. Tecta of two specins of Macroclieles showing lateral processes free (a) arid

region of the dorsal shield of Mucrocheles sp. cheles (b). partly fused (b).

Subfamily PARHOLASPINAE nov.

This subfamily shows far less dwersity in form than the Macrochelinae. All the known species are free-living in decaying vegetation. The sclerotization of the venter of the male is essentially the same in all the species examined, being provided with a holoventral shield formed by the fusion of the sternal, metasternal, podal, peritrematal and ventri-anal shields. The degree of fusion of the shields covering the venter of the female varies considerably as has already been noted (Fig. 1). This extreme variation in the degree of sclerotizatioii of the venter in closely allied groups of species questions the validity of adopting such characters in the supra-generic classification of the Mesostigmata.

The present concept of the evolution of the Parholaspinae is shown in Fig. 6. Berlese (1904 & 1918) has proposed four genera, Garnasholaspis, Holaspulus,

Parhohpis and Calholaspis, which have a straight peritreme and a fusion of the peritrematal and podal shields. These genera and three new genera described below are separated in the following key :

Key to the genera of the subfamily Parholaapinae

Females 1. Peritrematal shields fused with the ventri-anal shield

-. Peritrematal shield not fused with the ventri-anal shield (Fig.10) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.

(Fig. la) ............................................ 1.

CLASSIFICATION O F THE MACROCHELTDAE 353

2 . Cknital shield fuscd ~iosteriorly with thc vent ti-uiial shield (Fig. 12) ; dorsal seta oil chulicurn siinple (Pig. 16). . . . . . . . . .

- . Genital shield frce (Pig. 28) ; dorsal seta 011 cheliceril wodge-

2. Dorsal shield wit.h more than forty pairs of setae (Fig. 26) ; lihteral interscutal membrane with a thick covering of setae t-nsembling a pelage ; metaster~ial shields free (Fig. 2 8 ) . . . . . .

-. Doisal shield with thirty pairs of setae (Fig. 31) ; latreral interscutal membrane with normal setation ; nietasternal shields fused with the sternal and/or endopodal shields. . . . . .

4. Meta.sterna1 shields fused with t>he sternal ; ventri-anal shield with more than four pairs of preanal setae ; tarsus ii with stout spurs apically. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

-. Metasternal shields free (Fig. 42) ; ventri-anal with three or four pairs of preanal setae . . . . . . . . . . . . . . . . . . . . . .

5. Ventri-anal shield with three pairs of preamal setae ; tectum with lateral processes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

- . Ventri-anal shield wit,h four pairs of preanal setae ; tectuni

6. Dorsal shield with tweiity-nine pairs of long simple setae (Fig. 41) ; movable digit of the chelicera shorter than the corniculus, dorsal set& wedge-shaped ....................

-. Dorsal shield wit.h t~hirt,y pairs of setae of which twenty-nine pairs are hroadly lanceolate (Fig. 50) ; movable digit of the chelicera longer t'han the corniculus, dorsal seta simple. . . . . .

shaped (Fig. 34). . . . . . . . . . . . . . . . ....................

with lateral processes . . . . .

1.

2.

3.

4. -

Males

Dorsal shield witah t,weiity-iiine pairs of setae and surrounded by a wide expanse of membrane; movable digit of the chelicera shorter than the coriiiculus . . . . . . . . . . . . . . . . . . . . . Dorsal shield with t,hirty pairs of dorsal se1 a,c arid Rurrounded by a narrow band of inenibrane ; movable digit, of t,he chelicera slightly or considerably longer t,han t,he corniciilus Tectum without a median process (Pig. l a ) ; spermat,o- phoral process simple (Fig. 18). . . . . . . . . . . . . . . . . . . . . . . . . . Tectum produced into a mediaii process (Fig. 33) ; spermatophoral process with a ventral appendage (Fig. 36). . . . . . . . . . Tectuin wit.11 lateral processes (Fig. 61). . . . . . . . . . . . . . . . . . Tectum with a median process only.. ..................... Dorsal set& on chelicera wedge-shaped (Fig. 36) ; set,ae on dorsal shield simple or apatulat,e distally. . . . . . . . . . . . . . . . . . Dorsal set,a on chelicera simple (Fig. 63) : stttae on dorsal shield broadly lanceolate (Fig. 60). . . . . . . . . . . . . . . . . . . . . . .

Genus HOLASP UL. U S Red.

Nepcwholaspis gel). n.

C'ulholaspis Berl.

6.

Parh o1n.spultr.s gen. 11.

6.

Pnr1inlnspi.s Berl.

Dorsal shield entire with thirty pairs of setae of which twenty-nine pairs are spatulate distally. Extra-marginal set'ae spatulate. Female with sternal shield bearing three pairs of setae ; Genital shield with a pair of setae and fused posteriorly with t'he ventri-anal shield. Peri- trematal shield fused with the exopodal md ventri-aim1 shields. Male wit<h

metastenial shields free.

354 G. OWEN EVANS

holoventral shield : genital orifice praesternal. Both sexes with well developed prae-endopodal shields. Clorniculi and salivary styli long ; internal malae forked. Tectum with a denticnlated anterior margin. Chelicerae long and slender; movable digit longer than the corniculus. Dorsal seta on the chelicera simple : vent'ral seta long and pilose. Movable digit of the chelicera in the male with a strong spermatophoral process. Leg I in both sexes without ambulacra ; legs 11-IV with two claws and a lobate pulvillns. Femur and genu 11 in the male spurred.

Type : Holostaspis (Ho1a.spulus) tenuipes Berl.

Key to species 1. Genital aid ventri-anal shields evenly reticulated ; anterior

margiii of t#he geiiital shield ext>ending well beyond the posterior margin of the st,ernal shield. Male unknown.. . . .

- . Genital aid ventri-anal shields unevenly reticulated ; anterior margiu of t,he genital shield not, exteiiding beyond the posterior margin of the sternal shield. . . . . . . . . . . . . . . . . .

2. Sternal shield in the fpmale wit,h numerous closely set, puiictatioiis ; expulsory vesicles coiispicuous (Fig. 12). Spermatophoral process in t,he male considerably longer than the movable digit (Fig. 18). . . . . . . . . . . . . . . . . . . . . . . .

- . Sternal shield in the female without closely set puiirtations ; nxpulsory vesicles apparently absent (Fig. 21 ). Spermato- phoral process in t,he malt? shorter than t,he IIlOvahlt? digit, (Fig. 1 4 ) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Holusp,rtl~irs t m e d i r i sp. 11.

Ho1uspwltr.s vidii,lrn Re1.1.

3.

Holnspd?i..s trwuipe.? Bed.

HOLrlSPULUS I IENUIPES Berl. Holostrwpi.9 (Holuspid~ts) t e r w i p m Bcrlese, A. (1904) Hndia 1, 266. P(trltoIri.?pis pnch!ylcie.

Impsoiden Schweizer, J . (1932) Rw. xirisse Zoo/ . 39, 366, figs, S,I/PE. wo?:.

Ferntrle Dorsal shield strongly sclerotized and ornamented with punctations and fine

rebiculat'ions, especially in the posterior third. Dorsal setae with the exception of simple setae M I , spatulate distally. The distribution of the setae and pores are shown in Fig. 11. The extra-marginal setae lie on the strongly striated lateral interscutal membrane.

The sternal shield is covered with a network of ridges and closely set punctations ; its posterior margin is deeply emarginate ( Fig. 12). The three pairs of sternal setae are simple ; the third pair lying well inside the bases of the second. Each metasternal shield is free and provided with a simple seta and ' ' pore ". The genital shield is fused posteriorly with the ventri-anal shield. Its anterior hyaline portion is produced into a sharp point which almost reaches the posterior margin of the sternal shield. The endopodal shields in the region of coxae I1 are completely fused with the lateral margins of the sternal shield but in the region of coxae 111 and I V they are fragmenbed and not fused with the metasternal or genital shields. The ventri-anal shield is fused with the genital, podal and peritrematal shields. It is ornamented and bears four pairs of preanal setae in addition to t'he three setae normally associated with the anus. The position of the stigma and the form of

The tritosternum is well developed with the lacinae long and pilose.

All these setae are simple.

CLASSTFK’ATION OF THE MACROCHELIDAE 355

the peritremes are shown in the figure. The peritreniatal shield is fused posteriorly with the exopodal and ventri-anal shields.

An interesting feature of this species is the presence of a pair of expulsory vesicles posterior to coxae IV. In optical section each ‘‘ vesicle ” consists of a large gland from which issues a t’hick-walled duct terminating in a sacculus

12

Figs. 1 I-lf.-HoZuspuZz~s tcnctipes Berl. Female. 11. Dorsal shield. I?. T’ciit 1 . ~ 1 shields,

(Pig. 13). The sacculus is connected by a subsidiary duct to an opening in the ventral shield. Homologous structures have been reported in a number of gamasids and Michael considered them to be excretory organs on the basis of the ‘. yellowish oily fluid ” they contain (Michael, 1892). There is no definite evidence that they fuiiction as such.

The tubular gnathosoma is provided with the nornial four pairs of setae ventrally. The rostrals are considerably longer than the other three pairs (Fig. 14). The ventral groove is broad and provided with transverse rows of denticles. Each corniculus extends almost to the distal margin of the palp- femur. The internal malae are forked ; the internal process being considerably longer than the external. The tectum has a denticulate anterior margin (Fig. 15). All the setae on the free segments of the pedipalp with the exception of the three-pronged palptarsal seta, are simple. Both digits of the

356 G . OWEN EVANS

chelicerae are dentate ; the fixed being tridentate and the movable bidentate (Fig. 16). The movable digit (approx. 165 p in length) is considerably longer than the corniculus (approx. 125 p in length). The externo-ventral brush of setae and the pilose ventral seta are well developed. Leg I (approx. 640 p in length) lacks an arnbulacrum ; tarsus 1 (175 p) is about twice the length of

14

Figs. 13-1 9.--Ho/a,s~~?t/irs tenuipes Berl. 13. Expulsory vesicle. 14. Venter of the gnathosoma of the female. 15. Tectum. 16. Chelicera of the female. 17. Sickle-likc seta on leg 11 jn trhe female. 18. Chelicera o f the malo, 19. Leg I1 of the male,

CLASSIFICATION OF THE MACROCHELIDAE 367

tibia I (88 p). sickle shaped setae (Fig. 1).

Male

The four distal segments of legs 11-IV are provided with

The chaetotaxy of the dorsal shield is essentially the same as in the female. The venter is covered by a holoventral shield. In outline it resembles that

of the female. The region between coxae 11-IV is ornamented with a network of ridges (predominantly longitudinal) and punctations. The five pairs of setae in this region are simple. The reticulated ventri-anal shield bears four pairs of preanal setae.

The structure of the gnathosoma is similar to that in the female. The unidentate movable digit of the chelicera on the other hand, has an extremely long spermatophoral process, approximately 220 p in length (Fig. 18). The dentition of the fixed digit is shown in the figure. The movable digit is slightly longer than the corniculus (measuring 122 & 110 p, respectively).

Tibia I and tarsus I measure 176 and 77p respectively. Femur and genu I1 are spurred (Fig. 19).

Dimensions

The genital orifice is praesternal.

The chaetotaxy of the legs is essentially the same as in the female.

Female : length, 640-675 p ; breadth, 340-370 p. Male : length 570-585 p ; breadth, 295-305 p.

Habitat and Locality This species is known only from glasshouses in Europe (Italy, Berlese,

1904 and Switzerland, Schweizer, 1932). The above redescription of the species is based on specimens collected from soil and organic matter under Caryota mitis in the Aroid House at the Royal Botanic Gardens, Kew (Coll. P. N. Lawrence).

HOLASPULUS VIDUUS Bed. Holmpulus viduzcs Berlese, A . (1905) Redia 2, 164, Fig.

Berlese’s description of this species is short and does not mention many of the structures which are necessary to separate it from the new species described below. The key characters given above are based on the figure of the venter of the female, the only sex described, given by Berlese (1905). The length of the species is given as about 550p.

Habitat and Locality

ad Tjiompea in silvis collecta ”. It is known only from the type locality in Java : “ Nonulla vidi exempli

HOLASPULUS TWEEDIEI sp. n. Female

The ornamentation and chaetotaxy of the entire dorsal shield is similar to that in H . tenuipes (Fig. 20). The extra-marginal setae are spatulate distally and increase in length from the anterior to the posterior seta of the series. The lateral interscutal membrane is not as strongly striated as in tenuipes.

P.Z.S.L.-127 24

,368 0. OWEN EVANS

The sternal shield is strongly sclerotized and heavily ornamented with a The three

The The structure of the podal,

The gnathosoma is typical for the genus with the tectumdenticulated The movable digit of the chelicera

network of ridges which pass into punctations laterally (Fig. 21). pairs of sternal setae are simple. genital shield is fused with the ventri-anal shield. peritrematal and ventri-anal shieIds is shown in the figure.

along its entire anterior margin (Fig. 22).

The metasternal shields are free.

Figs. 20-24.-Holaspulzbs tweediei sp. n. 20. Dorsal shield of female. 21. Venter of female 22. Tectum of female. 23. Chelicera of female. 24. Chelicera of male.

CLASSIFICATION O F THE MACROCHELIDAE 359

is bidentate; the fixed quadridentate (Fig. 23). slightly longer than the corniculus (121 p).

Male The chaetotaxy and ornamentation of the dorsal shield is basically the same

as in the female. The holoventral shield is strongly sclerotized and ornamented with a network

of ridges and punctations in the sterniti-genital region. There are four pairs of simple preanal setae on the ventri-anal portion of the shield.

The movable digit of the chelicera is unidentate with the spermatophoral process considerably shorter than the digit (Fig. 24).

Leg I1 has the femur and genu spurred as in tenuipes.

Dimensions.

The fixed digit 130pis The dorsal seta is simple.

Tarsus I (approx. 154 p ) is about twice as long as tibia I (approx. 72p).

The tectum is the same as in the female.

The fixed digit is bidentate. The distal segments of leg I are missing in specimen examined.

Female : length, 520-535 p ; breadth, 270-275 p. Male : length, 480 p ; breadth 255 p.


Reservoir, Singapore by Mr. M. W. F. Tweedie on 3.x.1955. 1955 :

Four females and one male collected inside a rotting tree near the McRetchie Holotype female,

Allotype male, 1955 : 11 : 12 : 12 and Paratypes, 1955 : 11 : 22': 11 ; 11 : 12 : 13-15,

Genus TFUCHOLASPIS gen. n.

Dorsal shield entire with more than forty pairs of simple setae due to the fusion of the bases of a number of the setae normally situated on the interscutal membrane with the dorsal shield. Interscutal membrane with a dense covering of setae resembling a pelage. Sternal shield with three pairs of setae ; metasternal shields free. Genital shield free but surrounded by a perigenital shield formed by the fusion of the endopodal shields with the sternal and ventri- a n d shields. Peritrematal shield fused with the exopodals and the ventri-anal shield. Expulsory vesicles present. Gnathosoma normal for the Parholas- pinae. Tectum triangular. Chelicera chelate-dentate ; dorsal seta wedge- shaped. Leg I without ambulacra ; legs 11-IV each with two claws and a lobate pulvillus. Male unknown.

Type : Tricholaspis marginipilis sp. n.

TRICHOLASPIS MARGINIPILIS sp. n. Female

The reticulated dorsal shield is entire and bears, in addition to the thirty pairs of setae normally present in the Areolaspinae, about twenty-five pairs of simple setae on its lateral margin (Figs. 25 & 26). These setae are shorter than the other setae on the shield and originate from the lateral interscutal membrane which is densely beset with simple setae. Their fusion with the lateral margin of the shield is similar to the condition in Jordensia Oudms.

24*

360 (3. OWEN EVANS

The tritosternum has a narrow base and a pair of long pilose Iacinae. The sternal shield, heavily ornamented with a network of ridges, is completely fused with the endopodal shields. It bears three pairs of simple setae (Fig. 22). The metasternals are free. The genital shield, completely surrounded by a perigenital shield, has a pair of setae (Fig. 28). The ventri-anal shield is fused with the podal and peritrematal shields. Its surface is strongly reticulated and there are four pairs of preanal setae. The form of the peritreme is shown in the figure. There are a pair of expulsory vesicles postero-lateral to coxae IV.

25 Figs. 25-26.--TrichoZaspis marginipilis sp. n. Female. 25. Dorsal shield and lateral inter-

soda1 membrane. 26. Lateral margin of the dorsal shield.

The venter of the gnathosoma is normal for the family. The tectum is produced into a triangular process (Fig. 29). The digits of the chelicerae are short and strongly developed ; the movable digit (approx. 60 p) being considerably shorter than the corniculus ( 1 10 I / ) . The movable digit is bidentate ; the dentition of the fixed is shown in Fig. 30. The dorsal seta is wedge-shaped.

Leg I with tarsus (approx. 1751") is considerably longer than the tibia (approx. 115 p ) .

Dimensions

Tarsus I1 without spurs or stout spines.

Lengt'h, 630-650 E L ; breadth, 297-305 p.

CLASSlE’ICATION O F THE MACROCHELIDAE 361

Locality and Habitat Two females collected at the bottom of a tree near the McRetchie Reservoir,

Singapore by Mr. M. W. F. Tweedie (6 : ii : 1952). Holotype, 1995 : 11 : 22 : 16 ; Paratype, 1955 : 11 : 22 : 17.

met. sh.

end.sh.

30

Figs. 27-30.-Tricholmpis marginipilis sp . n. Female. 27. Venter. 28. Genital region. 29. Tectum. 30. Chelicere.

Abbreviations m in Fig. 1 8 (p.347)

Genus NEPARHOLASPIS gen. n. Dorsal shield with thirty pairs of simple or spatulatc setae. Lateral

Female with the metasternals Genital shield com-

Peritrematal shield fused with the

interscutal membrane with normal setation. fused with the sternal and/or the endopodal shields. pletely surrounded by a perigenital shield.

362 a. OWEN EVANS

podal and ventri-anal shields. Male with holoventral shield ; genital orifice praesternal. Gnathosoma normal for the ParJlolaspinae ; corniculus longer than the movable digit of the chelicera. Chelicerae chelate-dentate in both sexes ; movable digit in the male with a complex spermatophoral process. Leg I in both sexes without ambulacra ; femur, genu and tibia I1 spurred in the male. Species free-living in forest litter and humus.

Type : Neparholaspis sputhulutm sp. n.

Key to species Third pair of sternal setae in the female whip-like, considerably longer than the second pair; genital shield uniformly ornamented with reticulations. Male unknown. Nepurholrmpis crispus

Third pair of setae not whip-like, similar to second pair; genital shield weakly sculptured. ........................ Dorsal shield with strong reticulations, dorsal setae spatulate distally (Fig; 31) ; dorsal process of the spermatophoral process more than three times the length of the ventral process (Fig. 36). ..................................... Neparholaspis spathulatus

Dorsal shield weakly reticulated, dorsal setae, long, simple ; dorsal process of the spermatophoral process only slightly longer than the ventral process (Fig. 40). ..................

(Willmann).

2.

sp. n.

Neparholaspis Simplex sp. n.

NEPARHOLASPIS SPATHULATUS sp. n. Female

Dorsal shield, heavily ornamented with a network of ridges, bears thirty pairs of long setae which are distinctly spatulate distally (Fig. 31). The lateral interscutal membrane forms a narrow band laterally and posteriorly ; the extra-marginal setae are similar in form to those on the dorsal shield.

The venter is almost entirely covered by a strongly sclerotized shield formed by the fusion of the sternal, metasternal, podal, peritrematal and ventri-anal shields (Fig. 32). All the ventral setae are simple. The four pairs of preanal setae are distributed as in the figure.

The tectum has its anterior margin produced into a serrated triangular process (Fig. 33). The movable digit of the chelicera (Fig. 34) is bidentate and shorter than the corniculus (115 p & 140 p in length, respectively) ; the distal third of the fixed digit is multidentate.

Leg I (approx. 900 p in length) has the tibia (154 p ) shorter than the tarsus ( 220 p). Legs 11-IV have well developed ambulacra.

Mule The chaetotaxy of the dorsal shield is essentially the same as in the female.

The venter is almost entirely covered by a holoventral shield ornamented in the sterrial region with a network of ridges. There are no expulsory vesicles. The genital orifice is praesternal.

The tritosternum and prae-endopodal shields are well developed.

The sternal region is ornamented with a network of ridges.

There is a pair of large expulsory vesicles posterior to coxae IV. The gnathosoma is normal for the subfamily.

Tho dorsal seta is wedge-shaped (Fig, 35).

CLASSIFICATION OF THE MACROUHELIDAE 363

Figs. 31-37.-i?eparhoZaspis spathztZut?ts sp. XI. 31. Dorsd shield of the female. 32. Venter of female. 33. Tectum of the female. 34. Chelicera of the female. 36. Dorsal seta on chelicera. 36. Chelicera of the male. 37. Leg I1 of the male.

364 Q. OWEN EVANS

The tectum is of the same form as in the female. The unidentate movable digit of the chelicera is provided with a complex spermatophoral process ; the distal process being about 34 times the length of the ventral (Fig. 36). The dentition of the fixed digit is shown in the figure.

Leg I (approx. 825p) has the tibia (143p) shorter than the tarsus (180p). Leg I1 is armed with spurs as in Fig. 37.

Dimensions Female : length, 1050-11OOp ; breadth, 810-820 p. Male : length, 780-8OOp ; breadth, 710-715p.

Habitat and Locality Three females and two males from litter and topsoil in a forest at Sungei

Buloh, Selangor, Malaya ('2011. J. R. Audy, 1955). Holotype female, 1955 : 11 : 22 : 18 ; Allotype male, 1955 : 11 : 22 : 19 and Paratypes, 1955 : 11 : 22 : 20-22.

NEPARHOLASPIS SIMPLEX sp. n. Female

Dorsal shield with thirty pairs of simple setae distributed as in the preceding species.

The tritosternurn and prae-endopodal shields are well developed. The form and ornamentation of the ventral shields are shown in Fig. 38. The metasternal shields, fused with the endopodals, are separated from the sternal shield by an oblique fissure.

The gnathosoma is normal. The form of the tectum is shown in Fig. 39. The dentition of the chelicerae is essentially the same as in spathulatus.

Leg I (approx. 7 8 1 p ) has the tibia considerably shorter than the tarsus (132 & 187 p, respectively).

Male Dorsal shield and chaetotactic pattern essentially the same as in the

female. The venter is almost entirely covered by a holoventral shield as in the

preceding species. The sternal shield in the region of coxae I1 has an inverted V-shaped ridge.

The spermatophoral process on the movable digit of the chelicera is short with the ventral process approximately equal in length to the dorsal. The dentition of the digits is shown in Fig. 40.

Leg 1 (approx. 805 p ) with the relative lengths of the tibia and tarsus as in the female. Leg I1 is armed as in spathulatus but the axillary process is smaller.

Dimensions

All the setae on the venter are simple.

All the ventral setae are simple. The tectum is similar to that in the female.

Female : length, 920-950 p ; breadth, 620-630 p . Male : length, 814 p ; breadth, 605 p.

Two females and one male from forest litter and topsoil, Sungei Buloh, Habitat and Locality


Selangor, Malaya (coll. J. R. Audy, 1955). Allotype male, 1955 : 11 : 22 : 24 ; Paratype, 1955 : 11 : 22 : 25.

Holotype female, 1955 : 1 1 : 22 : 23 ;

./’

38 E’igs. 38-40.-Neparholaspis simplex sp. n. 38. Venter of female.

of male.

39 \l.**

39. Tectum. 40. Chelicers

NEPARHOLASPIS CRISPUS Willman comb. nov.

Parholaspis ( P ) crispus Willmann, C. (1940). Willmann, C . (1941).

2001. Anz. 130, 215. Parholaspis crispus Stud. Gebebiete Allg. Karstf. Berlin Biol. Ser. Nr. 8 , 27, figs.

This species is undoubtedly congeneric with N . spathulatus. It is known from a single female collected in a “ cave near Radna, Carniolia,” The characters used in the key to separate this species from spathulatus and simplex are taken from Willmann’s description and figures of the type. The dimensions are given as : length, 990 p, breadth, 930 p.

Species incertae sedis

PARHOLA8PIS CAELEBS Vitzthum

Parholaspis caelebs Vitzthum, H. G. (1926). Treubia 8, 35, figs.

The description and figures of the male of this species are insufficient to There is no doubt, however, that place it with certainty in its correct genus.

366 a. OWEN EVANS

it belongs to the Parholaspinae. In many ways it resembles the males of Neparholaspis and for this reason it is included, provisionally, in that genus. The type locality is stated to be in soil, Tjibodas, Java. It measures 695 p in length and 445 p in breadth.

Genus CALHOLASPIS Berl.

Cul7kolaspis Berleae, A. (1918).

Dorsal shield entire and bearing a number of long simple setae. female, the metasternal shields are fused with the sternal. shields are fused with the exopodals but not with the ventri-anal shield. latter is provided with six pairs of preanal setae. a long median spine.

Redia 13, 173

In the The peritrematal

The The tectum is produced into

The male is not known. Tarsi I1 has stout Hpurs distally.

Type : Calhlaspis swperbw Berl.

CALHOLASPIS SUPERBUi3 Berl.

Cnlholaspia superbus Berlese, A. (1918). Re& 13, 173.

This is t,he only species of the genus which has been described and has the characters given in the above definition of the genus.

Dimensions


Length, 1000 p ; breadth, 600p

In humus, Columbia, U.S.A.

Genus OAMASHOLASPIS Berl.

Bamasholaspis Berlese, A. (1904). Redia I, 265

Dorsal shield entire, incompletely covering the idiosoma and bearing twenty- nine pairs of long, simple setae. Lateral interscutal membrane extensive and provided with a moderate covering of simple setae. Female with strongly sclerotized sternal shield with three pairs of simple aetae ; metasternal shields free. Ventri-anal shield with four pairs of preanal setae and not fused with the podal-peritrematal shields. Male with holoventral shield. Gnathovoma with corniculi longer than the movable digits of the chelicerae ; movable digit in the male with a spermatophoral process. Dorsal seta wedge-shaped; ventral seta strongly developed. Tectum produced into a serrated median process. Free- living in decaying vegetable material.

Leg I longer than the dorsal shield and without ambulacrum.

Type : Holostaspis (Gamasholaspis) gamasoides Berl., 1904.


Key to species Females

1. Anterior margin of the ventri-anal shield deeply emarginate to accommodate posterior portion of the genital shield ; fused podal-peritrematal shields not flanking the ventri-anal shield; apical setae on palptarsus about one-third the greatest length of the segment. . . . . . . . . . . . . . . . . . . . Cfarnasholaspisgamasoides Berl.

-. Anterior margin of the ventri-anal shield truncated ; fused podal-peritrematal shields flanking the anterior region of the ventri-anal shield ; apical setae on palptarsus minute, subspinose, about one-sixth the greatest length of the segment ............................................ Gamasholuspis internzediris

sp. n.

CfAMASHOLASPIS GAMASOIDES Berl.

Holostmpis (Oarnasholaspis) gamasoides Berlese, A. (1904) Redia I, 265.

Female Its

surface is ornamented with, coarse punctetions. The lateral interscutal membrane is provided with a moderate covering of simple setae.

The tritosternum and prae-endopodal shields are well developed. The sternal shield extends to the middle of coxae I11 and is ornamented with a network of ridges laterally and coarse punctations medially (Fig. 42). The metasternal shields are small and free ; each is provided with a simple seta and a " pore ". The endopodal shields in the region of the sternal shield are fused with it but are fragmented in the region of coxae IV. The genital shield is long and narrow and extends beyond the level of the posterior margins of coxae IV. It bears a pair of simple setae. The anterior margin of the ventri-anal shield is deeply emarginate to receive the posterior part of the genital shield. It has four pairs of simple preanal setae in addition to the three setae normally associated with the anus. The metapodaIs are sma11. The fused podal-peritrematal shields are produced posterior to coxae IV. There is no fusion between these and the ventri-anal shield.

The corniculi and salivary styli are long and slender. The anterior margin of the tectum is produced into a serrated median process (Fig. 43). All the setae on the free segments of the palp are simple. The movable digit of the chelicera is bidentate and the fixed quadridentate (Fig. 44). The former (72p) is shorter than the corniculus (88 p).

The setae on Legs II- IV are simple.

Male The following description of the male is given by Berlese (1904) : " Mas

pedibus Becundi paris femure inferne calcare runcato auctis ; gem tuberculo minimo ; tibia tuberculo sat conspicuo ; tarso apice dentigero. Chela calcare

Dorsal shield with twenty-nine pairs of setae distributed as in Fig. 41.

The venter of the gnathosoma is normal for the sub-family.

The dorsal seta is wedged-shaped. Legs I were damaged in the specimen under study.

368 a. OWEN EVANS

recto, valido, retrosus directo, acuto, e tertia antica parte digiti mobilis exorto.”

Dimensions Female : Dorsal shield ; length, 638 p ; breadth, 308 p.

Idiosoma ; length, 760-790 p ; breadth, 400 p. Male : Idiosoma, length 700 p ; breadth, 390 p.

Figs. 41-44.-aamasholaspis gamasoides Berl. Female. 41. Dorsal shield. 42. Venter. 43. Tectum. 44. Chelicera.

Locality Specimens in the Berlese Collection were collected from decaying vegetable

material in Florence, Genoa and La Plata. The above redescription of the species is based on a single female extracted by a Berlese funnel from vegetable matter in decay (rotten twigs and black earth) from an excavation in a meadow in Avellaneda Park, Tucumhn City, Argentina (Coll. : Dr. P. Wygodzynsky, 22.iii.1953).

(:J,ASSIFICATION O F THE MACROCHELJDAE 369

GAMASHOLASPIS INTERMEDIUS sp. n.

Female The dorsal shield is enti,re and provided with twenty-nine pzirs of long.

simple setae distributed as in Fig. 45. The shield covers only part of the idiosoma being borded laterally and posteriorly by a wide band of striated membrane bearing a number of simple setae. The surface of the shield is reticulated and punctated.

Figs. 45-49.-GamashoZaspis intermediw sp. n. Female. 45. Dorsum of female. 46. Venter. 47. Tectum. 48. Chelioera. 49. Dorsal seta of chelicera.

370 Q. OWEN EVANS

The sternal shield is ornamented with a network of ridges and punctations , the three pairs of sternal setae are simple. The metasternal shields are small and free (Fig. 46) ; each has a seta and " pore ". As in the preceding species, the endopodals are fused with the sternal shield in the region of coxae I1 but not in the region of coxae IV. The form of the genital shield is shown in the figure. Unlike gamasoides, the anterior margin of the ventri-anal shield is truncate. It carries four pairs of preanal setae and its surface is covered with a network of punctate lines. In its anterior portion it lies in close proximity to the posterior extensions of the podal-peritrematal shields so that the genital shield appears to be enclosed in a perigenital shield. The metapodal shields are small.

The structure of the tectum and the dentition of the chelicera are shown in Figs. 47 & 48, respectively. The apical setae on the palptarsus are minute and subspinose.

Leg I is longer than the dorsal shield ; the tibia (1 10 p) being considerably shorter than the tarsus (210 p).

Dimensions

Locality and Habitat

Sumatra, collected by Mr. A. H. G. Alston, 29. iii.1954. 22 : 27 ; Paratypes 1955 : 11 : 22 : 28-29.

The gnathosoma is basically the same as in the preceding species.

The dorsal seta is wedge-shaped (Fig. 49).

All the setae on legs 11-IV are simple.

Length, 720-730 p ; breadth, 440-450 p.

Three females from forest litter at qarbuluan (1800 m.), south of Sidekalang, Holotype, 1955 : 11 :

Genus PARHOZAPSIS Berl.

Parholaspis Berlese, A. (1918) Redia 13, 174

Dorsal shield entire with thirty pairs of setae of which twenty-nine pairs are broadly lanceolate ; interscutal membrane narrow and with normal setation. Female with sternal shield bearing three pairs of setae ; metasternal shields free. Genital shield large and with a pair of setae. Ventri-anal shield not fused with podal-peritrematal or genital shield ; four pairs of preanal setae. Male with holoventral shield ; genital orifice praesternal. Chelicerae in both sexes chelate-dentate ; movable digit with spermatophoral process in the male ; dorsal seta simple. Corniculi shorter than the movable digits of the chelicerae. Tectum with denticulate anterior margin and an undivided median process. Known species are free-living decaying vegetable material.

Type : Parholaspis desertus Berl., 1918.

Key to species Females

1. Prae-endopodal shields present ; sternal shield punctated ; two pairs of preanal setae simple. . . . . . . . . . . . . . . . . . . . . . . .

- . Prae-endopodal shields absent ; sternal shield with network of ridges ; all preanal setse lanceolate . . . . . . . . . . . . . . . . . .

Pnrhot'aspis desertus Berl.

Parholmpis kewemis sp. n.

CLASSIFICATION OF T H E MACROCHELJDAE 371

PARHOLASPIS DESERTUS Berl.

Parholmpis desertus Berlese, A. (1918) Rediu 13, 174

Female

form of the dorsal setae are shown in Fig. 50. bearing lanceolate setae forms a narrow band laterally and posteriorly.

The surface of the dorsal shield is faintly reticulated ; the distribution and The lateral interscutal membrane

Figs. 60-53.-Parhola~~is desertus Berl. Female. 60. Dorsal shield. 61. Venter. 52. l’ectum. 53. Chelicera.

The tritosternum and prae-endopodal shields are well developed (Fig. 51). The sternal shield extends to the middle of coxae I11 ; its posterior margin

372 a. OWEN EVANS

is emarginate. The surface of the shield is ornamented with coarse punctations ; the sternal setae are simple. The metasternals are free and each bears a simple seta and a " pore ". The ornamented genital shield has a pair of setae and is not fused with the ventri-anal. The latter, triangular in outline (286 x 325 p) , is covered with a network of punctate lines and carries four pairs of preanal setae of which two pairs only are lanceolate. The podal-peritrematal shields are produced into triangular processes posterior to coxae IV. There are a pair of oval metapodal shields.

The tectum is denticulate with a long undivided median process (Fig. 52). Both digits of the chelicerae are strongly developed ; their dentition is shown in Fig. 53. The movable digit is considerably longer than the coriiiculus (respectively, 154 & 110p in length).

The gnathosoma is normal for the subfamily.

The dorsal seta is simple.

Figs. 54-55. Parholaspis desertus Bed. Male. 54. Chelicera. 55. Femur. genu and tibia of leg 11.

Leg I measures about 740 p in length and has the tibia (121 p) much shorter than the tarsus ( 1 5 4 ~ ) . Male

The structure of the chelicera and the armature of femur, genu and tibia of leg I1 are shown in Figs. 54 & 55, respectively. Dimensions

Female : length, 825 p ; breadth, 460 p. Male : length, 700p ; breadth, 4 5 0 ~ .

Habitat and Locality This species is previously known only from the male collected at Samarang,

Java (Berlese, 1918). The above description of the female is based on a single specimen collected by Mr. M. W. F. Tweedie from " roots of tree, Nee Soon Reservoir, Singapore, 20 : ii : 1952 ". Allotype, 1955 : 11 : 22 : 30. In addition, five females and two males were examined from forest litter and topsoil at Sungei Buloh, Selangor, Malaya (Coll. J. R. Audy).

Legs 11-IV are provided with lanceolate setae.

This sex is adequately described by Berlese (1918).

PARHOLASPIS KEWENSI sp. n. Female

Chaetotaxy of the dorsal shield essentially the same as in desertus. The tritosternum is well developed but prae-endopodal shields are absent.

CLASSIFICATION OF THE MACHROCHELIDAE 373

The sternal shield is strongly ornamented with ridges and shallow punctations (Fig. 56) ; the sternal setae are simple with the first pair about twice the length of the third. Each metasternal shield bears a simple seta and a " pore ". The genital shield is large and ornamented with ridges, especially in its anterior half'. The ventri-anal shield (363 x 460 p) has a network of punctate lines and punctations. The fused podal peritrematal shield extends posterior to coxae IV but is not fused with the ventri- anal shield.

All the preanal setae are lanceloate.

Figs. 56-58.-ParhoZasp~8 kewensis sp. n. Female. 56. Venter. 57. Tectum. 58. Chelicsra.

The structure of the tectum and the dentition of the chelicera are shown in Figs. 57 & 58, respectively. The movable digit of the chelicera ( 2 2 0 p ) is considerably longer than the corniculus (143 p).

Leg I, without an ambulacrum, has the tibia (143 p) shorter than the tarsus (176 p).

Dimensions

Legs 11-IV have well-developed ambulacra.

Length, 1010 p ; breadth, 660 p .

A single female collected from fallen fruits of Caryota mitis in the Aroid Holotype, 1955 :


House, Royal Botanic Gardens, Kew by Mr. P. N. Lawrence. 11 : 22 : 31.

Genus PARHOLASPULUS gen. n. Dorsal shield entire with thirty pairs of simple setae. Female with sternal

Genital shield shield bearing three pairs of setae ; metasternal shields free.

P.z.s.L.--Iz~ 25

374 G . OWEN EVANS

free ; ventri-anal shield with three pairs of preanal setae. Peritrematal shield partially fused with the exopodal shield but not fused with the ventri-anal. Prae-endopodal shields fragmented. Male with holoventral shield. Movable

59

Figs. 59-60.-ParholaspuZuus alstoni sp. 11. Female. 59. Dorsal shield. 60. Venter.

digit of the chelicera in both sexes longer than the corniculus ; dorsal seta simple. Spermatophoral process in the male with a ventral appendage. Tectum in both sexes with lateral processes in addition to a median process. Femur, genu, tibia and tarsus of leg I1 in the male spurred.

Type: Parholaspulus abtoni sp. n.

PARHOLASPULUS ALSTONI sp. n.

Female Dorsal shield with thirty pairs of simple setae and a number of " pores "

distributed as in Fig. 59. The shield is surrounded by a narrow band of membrane bearing the extra-marginal setae ; the posterior members of the extra-marginal series are setose.

The tritosternum has a narrow base with a pair of strongly pilose lacinae (Fig. 60). The prae-endopodal shields are fragmented and consist of fourteen

CLASSIFICATION O F THE MACROCHELIDBE 376

small shields situated as in the figure. The sternal shield bearing three pairs of setae, is ornamented with striae and punctate lines. The punctate lines form a hexagonal network in the median region of the shield. The metasternal shields are subcircular in outline arid each bears a seta and a " pore ". The wedge-shaped genital shield is provided with a pair of genital setae ; its surface is ornamented with punctute lines. Lying in close proximity to the posterior margin of the genital shield is a large ventri-anal shield weakly ornamented with a network of punctate lines. The three pairs of preanal setae are distributed as in the figure. The peritreniatal shield is partially €used with the exopodal shield and extenda posterior to coxa IV.

The corniculi are long and slender but are shorter than the movable digit of the chelicera. The tectum is characteristically shaped being produced 1st.crally into long, variously divided, processes (Fig. 61).

The gnatliosoma is normal for the subfamily.

The dentition of the chelicera is shown in Fig. 62.

Figs. 61-64.-Parholmpulus aLtoni sp. n. 61. Tectum of female. 62. Chelicera of female. 63. Chelicera of male. 64. Femur, genu and tibia of leg IV.

Leg I (approx. 625 p in length) has the tibia (88 p) shorter than the tarsus Tarsus I1 has ( 1 20 p).

a pair of strong spines apically. Legs 11-IV have well-developed claws and pulvilli.

25*

376 G. OWEN EVANS

Male

the female.

sternal.

processes. short and simple. (85 p ) and bears a stout spermatophoral process. a short seta-like ventral appendage (Fig. 63).

tarsus (154 p ) .

The chaetotaxy of the dorsal shield in the male is essentially the same a8 in

The venter is covered by a holoventral shield. The genital orifice is prae-

The tectum is basically the same as in the female having strong lateral The fixed digit of the chelicera is tridentate with the pilus dentilis

The movable digit (99p) is longer than the corniculus The latter, is provided with

Leg I: (approx. 5 7 2 ~ ) has the tibia ( 8 8 p ) considerably shorter than the

The lateral band of membrane is inconspicuous.

Femur, genu, tibia and tarsus I1 are spurrdd (Pig. 64).

Dimemiom Female : length, 550 p ; breadth, 275 p. Male : length 530p ; breadth, 2 5 5 ~ .


Kew (Coll. A. H. G. Alston, 1955). Allotype male : 1956 : 1 : 2 : 2.

A male and female from soil in No. 3 Fern House, Royal Botanic Gardens, Holotype female: 1956: 1 : 2 : 1 and

SUMMARY

1. The classification of the Macrochelidae is discussed. The family is divided into two subfamilies : Macrochelinae TrggBrdh (including Areolaspinae Trgggkdh) and Parholaspinae nov.

2. Keys are given to the genera (females only) of the Macrochelinae and to genera and species of the Parholaspinae.

3. Three new genera are proposed : Parhohpulw gen. n., type Parholaspulus alstoni sp. n. ; Tricholaspis

gen. n., type : Tricholaspis marginipilis sp. n., and Neparholaspis gen. n., Type : Neparholaspis spathulatm sp. n. 4. The following new species, in addition to the three species referred to

above, are described and figured : Holaspulus tweediei sp. n. (both sexes). Neparholaspis simplex sp. n. (both sexes). Qamasholaspis intermedim sp. n. (female only). Parholaspis kewensis sp. n. (female only).

5. Parholaspis pachylaelapsoides Schweizer 1932 is considered to be a synonym of HoJaspulw tenuipes Berl., 1904, and Parholaspis crispus Willmann 1940 is transferred to the genus Neparholaspis.

ACKNOWLEDGMENTS

I am extremely grateful to Mr. M. W. F. Tweedie (Singapore), Dr. J. R. Audy (Malaya), Dr. P. Wygodzynsky (Argentina), and Mr. A. H. G. Alston and Mr. P. N. Lawrence of the British Museum (Natural History) for collecting the material upon which the revision of the Parholaq$nae is based,

CLASSIFICATION OF THE MACROCHELLDAE 377

REFERENCES

BERLESE, A. (1904). BERLESE, A. (1905). BERLESE, A. (1910). BERLESE, A. (1918). EVANS, G. 0. (1956).

EVANS, G. 0. & BROWNING, E. (1956).

MICHAEL, A. D. (1892).

Acari nuovi. Manip. I. Acari nuovi Manip. IV. Redia 2, 1 5 6 1 7 6 , figs. Lista di nuove specie e nuovi generi di Acari. Centuria quarta di dcari nuovi.

Redia I, 235-252, 258-280.

Redia 6, 242-271. Redia 13, 115-190.

An introduction to the British Mesostigmata with keys to families and

British mites of the subfamily Macrochelinae Triig&rdh

On variations in the internal anatomy of the Gamasinae, especially in Trans. Linn. Soc. Lond.

genera.

( ~ a ~ ~ i n a - M a c r o c h e l i d ~ ) .

that of the genital organs and on their mode of coition. Zool. (2) 5, 281-324.

J . Linn. Soc. Lond. (2001.) (in the press).

Bull. Brit. Mus. (Nat. Hist.) (Zool.) 4, 1, 1-55.

OUDEWS, A. C. (1903). OUDEMANS, A. C. (1939). S~HWEXZER, J. in HOLZAPFEL, M. (1932). Die Gewachshausfeuna des Berner Botanischen

T R ~ ~ R D R , I. (1 949). Descriptions of two new genera of Mesostigmata (Acarina), Aspidilaelaps

T R ~ G ~ R D H , I. (1952). Acarina, collected by the Mangarevan expedition to South Eastern Brk.

Proc. ZOOZ. SOC. L o d . 118,

Rediu

New list of Dutch Acari 11. Acarologische Aanteekeningen CIX.

Tijdschr. Elat. 45, 1-52. Ent. Ber. 8, 272-280.

Gartens.

from Samoa and Protoholmpis from Peru.

Polynesia in 1934 by the Bernice P. Bishop Museum, Honolulu, Hawaii. 2001. (2) 4, 45-90.

TITRK, F. A. (1948). 82-125.

VALLE, A. (1953).

VITZTHUM, H. G. (1926). Malayische Acari. Treubia 8, 35. VITZTHUM, H. G. (1930). WILLMA", C. (1940).

WLLLMANN, C. (1941).

Rev. m i s s e 2001. 39, 365-370.

En.t. Medd. 25, 311-325.

Insecticolous Acari from Trinidad, B.W.I.

Revisione di generi e sottogeneri Berlesiani (Primo contributo). 38, 316-360.

Acarologische Beobachtungen. Zool. Jahrb. Syst. 59, 281-350. Neue Milben aus Hohlen der Balkanhalbinsel, gesamrnelt von Prof. Dr.

Die Acari der Hohlen der Balkanhalbinsel. Stud. Geebiete Allg. Karstf. K. Absolon, Briinn.

Berlin. Biol. Ser. Nr. 8 ; 27.

Zool. Am. 130, 215.

ON THE CLASSIFICATION OF THE FAMILY MACROCHELIDAE WITH PARTICULAR REFERENCE TO THE SUBFAMILY...

Documents

Transcript of ON THE CLASSIFICATION OF THE FAMILY MACROCHELIDAE WITH PARTICULAR REFERENCE TO THE SUBFAMILY...