Zoology Publications from Victoria University of Wellington—Nos. 66 and 67

Notes and Drawings from Type Material of Collembola

Notes And Drawings From Type Material Of Collembola

Publication of this paper is assised by a grant from the Victoria University of Wellington Publications Fund.

Abstract

This paper is primarily concerned with presenting illustrations prepared where possible from type material of 36 species of Collembola belonging to the families Onychiuridae and Hypogastruridae. Some relevant notes are also included on the taxonomy of the various species.

Introduction

Many earlier descriptions of Collembola were brief and not accompanied by the sufficiently detailed drawings now regarded as necessary in modern taxonomic studies.

Over the past twenty years I have been able to visit some of the world's important repositories for type material concerned with insect taxonomy. During the examination of the collections of Collembola in these institutions I prepared many drawings from type material or from specimens of species identified by their authors. In particular I visited the British museum of Natural History, London (for the sake of brevity designated in this paper as BMNH), the United States National Museum, Washington D.C. (USNM), the Museum of Comparative Zoology, Harvard University (MCZ), the Manchester Museum, England (MM), the Illinois State Natural History Survey, and others in both Europe and the U.S.A.

The idea behind the work was a world monograph on the Collembola. However, because the completion of such a monograph is as far away as ever, it occurred to me that some of my drawings could, if published, be beneficial to present and future workers on Collembola. Hence this paper, which is the first in a short series of papers that will include notes and figures of hitherto unillustrated or inadequately illustrated species of Collembola together with new synonymies.

Acknowledgements

In presenting this material I would like to thank the following: the Nuffield Foundation, England and the Carnegie Corporation, New York for travelling fellowships; my own University Council for refresher leave to enable me to undertake this work; and my friends and colleagues in many countries who assisted me. In particular I must mention: Mr N. Riley, Keeper of the Department of Entomology, BMNH, when I first went there, and Dr T. Clay; the late Dr W. D. Hinks, Director of MM, who located G. F. Carpenter's types for me; Dr J. F. G. Clarke, Curator of Insects, USNM, and the late Drs R. E. Snodgrass and C. J. Drake who gave me much help and encouragement while I was there; Dr P. J. Darlington at MCZ; and the late Dr H. B. Mills, Director, Illinois State Natural History Survey.

Species Covered in this Account

Tullbergia bisetosa Boerner

Tullbergia mixta Wahlgren

Tullbergia affinis Boerner

Paratullbergia callipygos Boerner

Spelaphorura willemi (Boerner)

Hymenaphorura cockleyi (Folsom)

Hymenaphorura subtenuis (Folsom)

Protaphorura armata aurantiaca (Ridley)

Protaphorura pseudarmatus (Folsom)

Onychiurus fimetarius (L)

Paronychiurus ramosus (Folsom)

Pseudonychiurus dentatus (Folsom)

Xenylla baconae Folsom

Xenylla cavernarum Jackson

Xenylla corticalis Boerner

Xenylla grisea Axelson

Xenylla longicauda Folsom

Xenylla maritima Tullberg

Xenylla mucronata Axelson

Xenylla rhodesiensis Womersley

Xenylla schillei Boerner

Xenylla welchi Folsom

Hypogastrura armata communis (Folsom)

Hypogastrura cavicola (Boerner)

Hypogastrura gracilis (Folsom)

Hypogastrura humi (Folsom)

Hypogastrura macgillivrayi (Folsom)

Hypogastrura myrmecophila Womersley

Hypogastrura nivicola (Fitch)

Hypogastrura pseudarmata (Folsom)

Hypogastrura purpurascens galiciana (Boerner)

Hypogastrura viatica (Tullberg)

Hypogastrura (Cyclograna) boletivorus (Packard)

Schaefferia variabilis Christiansen

Proxenyllodes reticulatus Boerner

Triacanthella alba Carpenter

Figs. 1-8 Tullbergia bisetosa Boerner, drawn from hypotypes in B.P. Bishop Museum, Honolulu.

Figs. 1-8 Tullbergia bisetosa Boerner, drawn from hypotypes in B.P. Bishop Museum, Honolulu.

Fig. 1 Ant IV and apex Ant III × 1250
Fig. 2 anal spine × 1250
Fig. 3 genital opening of male × 1250
Fig. 4 apex of PAO beside cuticular granules × 2500
Fig. 5 section of PAO × 2000
Fig. 6 female genital aperture × 1250
Fig. 7 hind foot × 1250
Fig. 8 Abd VI × 320

Tullbergia mixta Wahlgren, 1906
Figs. 9-15

This species was recorded by Enderlein (1909) and by Gressitt and Weber (1959) from Graham Land and the South Shetland Islands. Several specimens were also collected by T. S. and R. E. Leech in 1961 from Admiralty Bay F.I.D.S. Base, Deception Island and from Greenwich Island, South Shetland Islands. I have identified this species recently in New Zealand at Riwaka, near Nelson. This species also, has not previously been illustrated. The figures given here are from specimens from Admiralty Bay. These specimens mounted on slides are deposited as hypotypes in the B.P. Bishop Museum, Honolulu.

Notes: Clothing on Ant IV includes stout curved blunt-nosed sensory setae, three short slender tapering apical sensory setae, 6-8 longer tapering somewhat blunt-nosed sensory setae, a single short subapical sense-rod in a pit, an apical trilobed sensory vesicle, and numerous long setae especially around the posterior border of the segment (Fig. 12). SO Ant III with four bent sense clubs and two very small sense-rods in pits with four guard setae, which is a much more complicated sense organ than originally described (Fig. 13); Abd VI anterior to the anal spines, with two parallel rows of five setae each (Fig. 9); Figs. 10 and 11 show the setae of the genital region. The claw (Fig. 15) bears a number of apical tibiotarsal setae and two basal setae, and is granulated basally. Some feet show an enlarged granule where the unguiculus should arise but this does not appear to be a normal feature. The PAO has 48-52 vesicles as in Fig. 14.

The body clothing is moderate with both short and long simple setae.

PSO one on each antennal base, one on each hind margin of head ThI 0+0, ThII 0+0, ThIII 1+1, AbdI 0+0, AbdII 0+0, Abd III 0+0, AbdIV 1+1, AbdV 1+1, AbdVI 0+0.

Figs. 9-15 Tullbergia mixta Wahlgren, drawn from hypotypes deposited with the B.P. Bishop Museum, Honolulu.

Figs. 9-15 Tullbergia mixta Wahlgren, drawn from hypotypes deposited with the B.P. Bishop Museum, Honolulu.

Fig. 9 Abd VI with anal spines and associated dorsal setae × 320
Fig. 10 genital aperture of male × 800
Fig. 11 genital aperture of female × 512
Fig. 12 Ant IV × 800
Fig. 13 SO Ant III × 1000
Fig. 14 PAO × 1280
Fig. 15 middle foot × 1000

Paratullbergia callipygos (Boerner, 1903)
Tullbergia callipygos Boerner, 1903
Figs. 20-23

Included in the collection of Boerner's material at the BMNH is a tube of specimens identified as T. callipygos but without any type citation. From these, which are most likely the type series, I selected one specimen, now mounted on a slide, which I designate as lectotype and from which Figs. 20-23 were prepared.

Notes: Clothing sparse of short simple setae, longer posteriorly. PSO —ant base, posterior margin of head, ThII-Abd V each 1+1. PAO of 60-70 vesicles. Ant IV with 4-5 stout curved sense rods, and several long plain setae but I could not see the apical sensory knob mentioned by Boerner. SO Ant III as in Fig. 20. Papillae of anal spines more coarsely granulated than those of the body. Abd III dorsally with four low cuticular ridges less pronounced than in most species of the genus Paratullbergia.

Hymenaphorura subtenuis (Folsom, 1917)
Onychiurus subtenuis Folsom, 1917
Figs. 32-36

These figures are drawn from a specimen labelled cotype USNM No. 20764.

Notes: The arrangement of the setae of Abd VI, on my interpretation, differs slightly from that of Folsom. The claw is slightly granulated basally and the whip-like extension of the unguiculus is relatively shorter, the unguiculus being more lance-like.

Protaphorura armata aurantiaca (Ridley, 1880) nov. comb.
Protaphorura armata denticulata (Handschin, 1924)
Onychiurus subaequalis Bagnall, 1937
Onychiurus flavescens Bagnall, 1935 syn. nov.
Lipura aurantiaca Ridley, 1880

Ridley's holotype is preserved in BMNH. After examining this specimen I find that it is identical with the subspecies denticulata Handschin, 1924, and therefore Ridley's name aurantiaca takes priority over the name denticulata for this subspecies.

The type of L. aurantiaca was overlooked when my paper on the Onychiuridae was published in 1959. The hind foot which is the distinguishing structure is shown on p.135 of that paper, Fig. 12. This species is also discussed in the footnote on p.164 of my "Index to the Collembola", 1964. As Bagnall's species flavescens has a denticulate claw it is a synonym of this subspecies, and my placing of it as a synonym of P. armata in my "Index", p.166, was incorrect.

Onychiurus fimetarius (L, 1758)
Onychiurus pseudofimetarius Folsom, 1917
Figs 41-43

In my "Index to the Collembola", (p.180) I recorded Folsom's O. pseudofimetarius as a synonym of O. fimetarius L. The figures drawn here are from a specimen labelled cotype of O. pseudofimetarius, USNM No. 20762. Folsom separated his species from fimetarius L. by the extra papilla of SO Ant III and the ventral PSO of the head. Stach (1954) records O. fimetarius with five papillae in SO Ant III and so far as my knowledge goes of this species this is normal structure for this organ. The PAO drawn has 18 vesicles; Stach records 18-24. Two PSO on the ventral surface of the head is normal for fimetarius. It appears, therefore, that Folsom's species is a junior synonym of O. fimetarius L.

Paronychiurus ramosus (Folsom, 1917)
Onychiurus ramosus Folsom, 1917
Figs. 44-49

The figures given here were drawn from a specimen labelled cotype in rather poor condition, USNM No. 20761.

Notes: The granulated sense clubs and smooth sense rods of the SO Ant III (Figs. 44 and 45) indicate that this species should be included in the genus Paronychiurus Bagnall, 1948. Relative to the segment Ant III this sense organ is very small. Fig. 47 indicates the setae associated with the anal spines on Abd VI.

Pseudonychiurus dentatus (Folsom, 1902)
Onychiurus dentatus Folsom, 1902
Figs. 50-54

A specimen labelled cotype USNM No. 5436 was used to prepare these drawings.

Notes: The PAO (Fig. 53) is more irregular in outline than suggested by Folsom. The claw as well as having a pair of external lateral winglike teeth and a pair of external distal teeth also has an inner tooth situated on an inner lateral "blade" or keel-like structure separate from the main inner keel of the claw. This "blade" swells distally into a kind of hummock or basal sheath (Fig. 51). The bases of both the claw and unguiculus are granulated. The papillae of the anal spines have granules much smaller than those of the surrounding cuticle (Fig. 54).

PSO as follows: Ant base 2+2, hind margin of head 1+1, ThI 0+0, ThII 2+2, ThIII 1+1, Abds I-V 2+2 each.

Xenylla cavernarum Jackson, 1927
Figs. 61-65

The holotype and some paratypes are in the BMNH and these drawings have been made from a paratype which is better preserved than the holotype. These are supplementary to those given by Jackson (1927, p. 486) in his original description.

Xenylla corticalis Boerner, 1901
Figs. 66-70

A specimen from the Boerner collection in BMNH was used for these drawings which supplement those of Boerner's original description and later of Axelson (1907, pl.V, Figs. 12-14).

Notes: The structural detail of the furcula indicates a spatulate shape with a distinct internal lamella for the mucro which is not distinctly separated from the dens. Ant IV has four subapical curved sense rods, an apical sensory knob in a pit and numerous simple setae.

Xenylla maritima Tullberg, 1869
Figs. 83-84

These two figures drawn from a specimen in the BNMH identified by Boerner supplement those of Stach (1949 pl.xxiii).

Xenylla mucronata Axelson, 1903
Figs. 85-89

Axelson's (1912, pls. iv and v) figures of this species deal with the furcula and anal spines. Specimens from BMNH Boerner's collection have been used for the figures included here and these add considerably more detail for the identification of this species.

Notes: Sensory structures of Ant IV are as drawn in Fig. 85. The SO Ant III (Fig. 87) is similar to that of X. longicauda Folsom with the two inner small sense rods completely hidden behind a high cuticular fold. The two tenent hairs of each foot of equal length (Fig. 86); papillae of anal spines contiguous; mucro incompletely separated from dens which bears two simple setae only on posterior face (Fig. 89). The species X. longicauda and X. mucronata are very similar differing mainly in the shape of the mucro and the extent of its lamella, and by the form and degree of separation of the anal spines and their papillae.

Xenylla schillei Boerner, 1903
Figs. 86-100

A specimen labelled cotype in the Boerner collection BMNH was used to prepare the accompanying figures. Drawings of the foot and furcula were included by Stach (1951, p.68).

Notes: The two tenent hairs of each foot are at different levels on opposite sides of each tibiotarsus (Fig. 99). The two small sense clubs of SO Ant III are almost completely hidden by the large cuticular fold (Fig. 98).

Hypogastrura myrmecophila Womersley, 1926
Figs. 140-143

A specimen labelled cotype in BMNH was used to prepare these drawings which are additional to those given by Womersley (1926, p.156).

Notes: PAO (Fig. 142) has five peripheral lobes, not four, the unguiculus has a broad inner lamellae (Fig. 140).

Hypogastrura nivicola (Fitch, 1847)
Podura nivicola Fitch, 1847
Achorutes socialis Uzel, 1890
Achorutes nivicola Folsom, 1902
Figs. 144-150

I have examined a specimen in the Boerner collection at the BMNH labelled A. socialis by Boerner, and also specimens labelled as cotypes of P. nivicola Fitch which are in the USNM (3 specimens on slide No. 5216). Considering the age of Fitch's specimens they were in a remarkably good state of preservation and figures 146-150 of this paper were drawn from them. Figs. 143-145 are from Boerner's specimens in BMNH. From these examinations it is clear that the synonomy suggested by Schott (1893) and adopted in my "Index" in 1964 is correct. H. nivicola was redescribed and illustrated by Stach (1949, pp. 71-77, pl.iv). Stach's figures of the species agree very clearly with those given here from Fitch's type material. The species would seem to be very widely distributed over the Northern Hemisphere.

Since these cotypes of Fitch were available in the USNM for Folsom to study, the suggestion made by Folsom (1916, p.485) that Fitch's name be suppressed in favour of socialis is quite out of order.

Notes: "Nebenhocker-like" structure beside PAO (Fig. 149).

Hypogastrura pseudarmata (Folsom, 1916)
Achorutes pseudarmatus Folsom, 1916
Figs. 151-157

Three specimens labelled cotypes of this species are mounted on slide No. 19901 USNM and the drawings of Figs. 151-157 were made from these specimens.

Notes: The peculiar arrangement of the PAO with the roundish "Nebenhocker" of Folsom adjacent to the nearest ocellus is shown in Figs. 152 and 153. SO Ant III has a slightly different displacement of the parts (Fig. 156) from that given by Folsom in his Fig. 97; each tibiotarsus has a single clavate tenent hair and a long subequal stout seta (Fig. 155); claw with a strong distal outer tooth as well as the outer basal tooth described by Folsom (Fig. 155); the papillae of the anal spines (Fig. 157) not quite contiguous as suggested in the original description; seven short simple setae on posterior face of dens (Fig. 154).

Hypogastrura (Cyclograna) boletivora (Packard, 1873)
Achorutes boletivorus Packard, 1873
Achorutes marmoratus Packard, 1873
Achorutes pratorum Packard, 1873
Achorutes armatus (Nicolet) Folsom, 1916
Hypogastrura (Cyclograna) vulgaris Yosii, 1960
Cyclograna vulgaris Yosii, 1960
Hypogastrura vulgaris Salmon, 1964
Figs. 164-171

Dr K. A. Christiansen has in his collection at Grinell, Iowa, a paratype of Cyclograna vulgaris sent him by Dr Yosii. I have studied this specimen and also Folsom's specimens labelled cotypes at USNM and the material left by Packard and preserved in MCZ.

Folsom (1916) who studied Packard's material considered that the three species described by Packard (1873) were all identical with A. armatus of Folsom. After studying all this material I agree with the suggestions made by Yosii in 1960, p.264, that Folsom's A. armatus and Yosii's C. vulgaris are the same species. This means that the correct name for this assemblage will be H. (Cyclograna) boletivorus (Packard), being the first species described by Packard (page priority) in his 1873 paper. However, the peculiar spines of the dens, the lamellae of the mucro and the odd PAO seem to call for the retention of Yosii's subgenus Cyclograna for this and the several related species described by Yosii. The occurrence of the eversible sac between Ants III and IV, the general body form and chaetotaxy all indicate that this species is a Hypogastruran form of Collembola. This view is further strengthened by the mouth parts; the mandible is present with a distinct molar area and the maxilla head has six apical teeth.

Figs. 164-171 are from the paratype of Cyclograna vulgaris Yosii and supplement those already given by Folsom (1916) and Yosii (1960).

Notes: Dens and mucro (Figs. 169-170) with their peculiar spines and lamellate mucro rather similar to the condition found in the genus Clavontella; furcula is reduced and very small, the dens and mucro being sub-equal in length to the hind claw; PAO is rather Folsomia-like on the surface but divides into three lobes as the microscope is focussed deeper into the cuticular groove in which the PAO lies.

Hypogastrura viatica (Tullberg, 1872)
Achorutes murorum (Bourlet, 1843) (Tullberg, 1869)
Achorutes viaticus Tullberg, 1872
Achorutus titahiensis Salmon, 1943
Achorutus subviaticus Bagnall, 1947
Achorutus pseudoviaticus Bagnall, 1947

Tullberg's (1872, pl.10, Figs. 7-20) original description of this species is beautifully illustrated, and further illustrations were given by Stach (1954, pls. iii, X) when he redescribed this species. Bagnall (1941, p.220) stated that the true viaticus of Tullberg had a tenaculum with four barbs and he set up two new closely related British species (A. subviaticus and A. pseudoviaticus) for forms he had discovered with only three barbs to each ramus. In 1943 (p.376) I described a new species A. titahiensis from New Zealand with three barbs to each ramus. Bagnall also stated that the papillae of the anal spines were contiguous basally whereas in his two new species these structures were more or less separated. In A. titahiensis these structures are variable and the anal papillae may be either continguous or separated. Stach (1949) in his redescription of A. viaticus records the rami of the tenacalum as each having three barbs and the anal papillae as being contiguous basally. He suggests that Bagnall's two species are synonyms of A. viaticus, a synonomy with which I would now agree.

I have examined a large series of specimens of A. viaticus at BMNH from various parts of the world and find that the papillae of the anal spines may be touching at their bases or close together but not widely-separated. The variations may occur amongst individuals from the same locality. The anal spines themselves vary considerably in length from very short (about 1/6 of hind claw) to quite long (about half length of hind claw). They may be straight but are normally slightly curved. One specimen from Dublin has two spines on one side of the body arranged one behind the other, and only one spine on the other side.

The tenaculum normally has only three barbs to each ramus but occasional specimens do occur with four barbs to each ramus. One specimen examined from Bear Island had four barbs on one ramus and three on the other.

Re-examination of my A. titahiensis type material convinces me that this is simply a further, perhaps extreme, variation of A. viaticus Tullberg.

In the BMNH collections there are some specimens labelled A. murorum and presented to the museum by Lubbock in 1884 from material sent to him by Tullberg. After examining these I consider them identical with A. viaticus Tullberg, 1872.

Proxenyllodes reticulatus (Boerner, 1909)
Hypogastrura reticulata Boerner, 1909
Figs 183-189

The type of this species is in the Boerner collection at BMNH. No figures were given with Boerner's original description but Yosii (1960) gives excellent figures of the chaetotaxy and some other structures of this species drawn from further specimens he collected in Central Japan.

Notes: The coarse granulations of the cuticle place this species, in my view, in the genus Proxynellodes Denis, 1926. This I consider preferable to retaining it as a "group" in Hypogastrura as suggested by Yosii. Ant IV (Fig. 183) with two apical protrusible sensory sacs, 7-9 short, bladderlike sensory rods and many simple setae. PAO (Fig. 186) with four contiguous lobes arranged more or less as a cross; claw and unguiculus both granulate basally, the latter with a granulate basal finger-like process lying alongside the base of the inner lamella (Fig. 184).

Triacanthella alba Carpenter, 1909
Figs. 190-198

Specimens labelled paratypes of this subantarctic species are in BMNH and these drawings supplement those included in the original description. Two specimens labelled cotypes on slides in the MM were also examined but both are badly damaged.

Notes: The claw has an internal tooth and a pair of outer distal lateral teeth (Figs. 193 and 194) but no tenent hairs; clothing of both short and long simple setae with some longer serrated setae especially around the posterior region.

Figs. 172-182 Schaefferia variabilis Christiansen, drawn from holotype and cotypes USNM.

Figs. 172-182 Schaefferia variabilis Christiansen, drawn from holotype and cotypes USNM.

Fig. 172 apex Ant IV × 1000 from cotype USNM
Fig. 173 PAO and adjacent ocellus × 1000 from cotype USNM
Fig. 174 PAO, adjacent ocellus and setae × 1000 from holotype USNM
Fig. 175 SO Ant III × 1000 from cotype USNM
Fig. 176 head of maxilla × 1000 from cotype USNM
Fig. 177 mandible × 1000 from holotype USNM
Fig. 178 lateral view of mucro × 1000 from holotyp USNM
Fig. 178 lateral view of mucro × 1000 from holotype USNM
Fig. 180 mucrodens × 600 from cotype USNM
Fig. 181 hind foot × 600 from cotype USNM
Fig. 182 anal spine and papilla from side × 600 from holotype USNM

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1930: Notes on some new and rare British Collembola. Ent. Mon. Mag. , 66, pp. 33-41.

Yossi, R., 1954: Die Kulturpflanzenschadigen Collembolen Japans. Oyo-Kontyu , 10 (2), pp. 137-141.

1960: Studies on the Collembola genus Hypogastrura . Amer. Midl. Nat. , 64 (2), pp. 257-281.

Professor J. T. Salmon, D.Sc., F.R.S.N.Z.
Zoology Department, Victoria University of Wellington, Private Bag, Wellington, New Zealand.

* In my "Index" on p.82 this paper is recorded as being published in 1929. This date is incorrect and should be amended to 1926.