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.
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.
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.
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 Accountpage 3
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 bisetosa Boerner, 1903
This species has been reported from a number of collections in different subantarctic regions but no illustrations of it have been published.
I have recently identified specimens of T. bisetosa from collections made by Dr J. L. Gressitt and by K. Watson from Macquarie Island. The accompanying illustrations are from specimens collected by Dr Gressitt on the moss Azorella, at the north end of Macquarie. These specimens are mounted on slides and deposited as hypotypes in the B.P. Bishop Museum, Honolulu.
Notes: Upper surface of Ant IV with very long simple setae and fine curved sense rods one of which is much longer than the others (Fig. 1); lower surface with many short simple setae. S.O.Ant III has three guard setae on the main organ, not two as described by Boerner, and two at the third separate sense rod. Details of the PAO and genital apertures as shown in Figs. 4, 5, and 3 respectively.
Tullbergia mixta Wahlgren, 1906
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.page 5
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
Tullbergia affinis Boerner, 1903
These drawings were made from the holotype in the BMNH. They agree very closely with the figures of this species drawn by Axelson (1906) and later reproduced by Womersley (1930, p.37). Stach 1951, p.15 and 1954, pl.26) figured this species but his Fig. 1 of the PAO appears to have been taken from a species other than affinis. Gisin (1960, pp.19, 154) also included two figures dealing with this species.
Notes: Length 1.2 mm. Ant IV with apical knob and six sub-apical curved sense rods. PAO in a deep cuticular groove, claw with short broad seta to each side, and very small vestigial unguiculus. Anal spines strongly curved, on papillae almost half as long as the spines. Clothing of short setae.
Paratullbergia callipygos (Boerner, 1903)
Tullbergia callipygos Boerner, 1903
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.
Spelaphorura willemi (Boerner, 1901)
Aphorura willemi Boerner, 1901
Onychiurus willemi Boerner, 1901, sensu Salmon, 1964
The holotype is in BMNH but is in poor condition, with the antennae missing. Boerner (1901, p.334), when he described this species included figures of the SO Ant III and PAO, and additional figures of the foot and anal spines are given here. From the drawings of the sense organs made by Boerner it is clear that this species does not belong to Onychiurus (Aphorura) and should be transferred to the genus Spelaphorura Bagnall, 1948.page 7
Figs. 16-19 Tullbergia affinis Boerner, drawn from holotype in BMNH.
Fig. 16 SO Ant III × 1000
Fig. 17 PAO × 1000
Fig. 18 hind foot × 400
Fig. 19 anal spines × 320
Figs. 20-23 Paratullbergia callipygos Boerner, drawn from lectotype in BMNH.
Fig. 20 SO Ant III × 1000
Fig. 21 PAO × 1250
Fig. 22 hind foot × 400
Fig. 23 anal spine × 320
Figs. 24-25 Spelaphorura willemi (Boerner), drawn from holotype in BMNH
Fig. 24 apex of Abd VI showing anal spines and associated setae × 500
Fig. 25 hind foot × 500
Hymenaphorura cockleyi (Folsom, 1908)
Aphorura cockleyi Folsom, 1908
Onychiurus similis Folsom, 1917
Syntypes labelled cotypes of this species are preserved in the USNM and the Illinois State Nat. Hist. Survey at Urbana. I examined the cotypes of A. cockleyi and O. similis in the former institution and drew the figures given here from a slide of mounted specimens (No. 12033) which I now designate as lectotype of H. cockleyi.
Notes: SO Ant III is very prominent at the apex of Ant III, and large in relation to the segment. The PAO is as in Fig. 29 and differs from that shown in Folsom (1917, Fig. 19). The setae of Abd VI in relation to the anal spines are as in Fig. 30. The bases of the claw and of the unguiculus are coarsely granulated, and the cuticular granules in general are relatively large.
The syntype of Folsom's Onychiurus similis in the USNM (No. 20763) is in poor condition but the setae of Abd VI are exactly as in cockleyi and the anal spines on their papillae are not continguous but separated as in cockleyi. The PAO, SO Ant III and foot are all almost identical with those structures in cockleyi and I am convinced that the two species described by Folsom are conspecific. O. similis, therefore, becomes a junior synonym of H. cockleyi.
Hymenaphorura subtenuis (Folsom, 1917)
Onychiurus subtenuis Folsom, 1917
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.page 9
Figs. 26-31 Hymenaphorura cockleyi (Folsom), drawn from lectotype in USNM.
Fig. 26 SO Ant III from side showing size in relation to Ant III × 420
Fig. 27 SO Ant III × 1000
Fig. 28 hind foot × 1000
Fig. 29 PAO × 1500
Fig. 30 Abd VI with anal spines and dorsal setae × 420
Fig. 31 anal spine and papilla from side × 1000
Figs. 32-36 Hymenaphorura subtenuis (Folsom), drawn from cotype in USNM.
Fig. 32 hind foot × 1000
Fig. 33 Abd VI with anal spines and dorsal setae × 420
Fig. 34 SO Ant III × 1500
Fig. 35 PAO and associated setae × 1000
Fig. 36 anal spine from side × 420
Protaphorura pseudarmatus (Folsom, 1917)
Onychiurus pseudarmatus Folsom, 1917
These figures are drawn from a specimen labelled cotype. USNM No. 20760.
Notes: The furcula is represented by a cuticular fold or ridge but this is very rudimentary and lacks any furcula-like structural differentiation. Folsom's species originally described as belonging to the genus Onychiurus clearly belongs in Protaphorura.
Onychiurus fimetarius (L, 1758)
Onychiurus pseudofimetarius Folsom, 1917
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
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
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.page 12
Figs. 37-40 Protaphorura pseudarmatus (Folsom), drawn from cotype in USNM.
Fig. 37 PAO × 1000
Fig. 38 SO Ant III, sectional view from side with end papilla removed × 1000
Fig. 39 anal spine × 1000
Fig. 40 fore foot × 1000
Figs. 41-43 Onychiurus fimetarius (L), drawn from cotype of Onychiurus pseudofimetarius Folsom in USNM.
Fig. 41 SO Ant III × 1000
Fig. 42 PAO × 1500
Fig. 43 forefoot × 1000
Figs. 44-49 Paronychiurus ramosus (Folsom), drawn from cotype in USNM.
Fig. 44 SO Ant III × 1500
Fig. 45 SO Ant III side view × 1500
Fig. 46 PAO × 1500
Fig. 47 Abd VI with anal spines and setae angled view × 530
Fig. 48 anal spine from side × 1000
Fig. 49 hind foot × 1000
Figs. 50-54 Pseudonychiurus dentatus (Folsom), drawn from cotype in USNM.
Fig. 50 SO Ant III from side × 1000
Fig. 51 middle foot × 600
Fig. 52 PAO, deep appearance at bottom of organ × 1000
Fig. 53 PAO surface appearance × 1000
Fig. 54 anal spine from side × 1000
Xenylla baconae Folsom, 1916
I have examined specimens labelled cotypes in USNM and in MCZ. The drawings here are from USNM specimen No. 19903.
Notes: Fig. 55, Folsom's drawing of the SO Ant III (Fig. 143 in his 1916 paper) shows one of the large sense rods as a seta. The apex of Ant IV (Fig. 56) has an exsertile sensory knob and sensory rods as shown; tenaculum (Fig. 59) with three barbs; each anal spine with an associated anterior-lateral seta (Fig. 60): the claws with one long simple ventral seta, though Folsom mentions two; and the dental hooks (Fig. 58) which are very well developed. I could not detect the central incision shown by Folsom in the mucro of this species.
Xenylla cavernarum Jackson, 1927
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
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 grisea Axelson, 1900
These figures of the hind foot and mucro, drawn from a specimen labelled cotype in BMNH show more detail than those of Axelson (1912) and later of Stach (1949, pl.22).page 15
Figs. 55-60 Xenylla baconae Folsom, drawn from cotype in USNM.
Fig. 55 SO Ant III × 1000
Fig. 56 apex Ant IV × 1000
Fig. 57 hind foot × 1000
Fig. 58 mucrodens × 1000
Fig. 59 tenaculum × 1000
Fig. 60 anal spine and associated seta from side × 1000
Figs. 61-65 Xenylla cavernarum Jackson, drawn from paratype in BMNH.
Fig. 61 Ant IV and apex Ant III showing SO Ant III × 1000
Fig. 62 mucrodens × 400
Fig. 63 ocelli and associated seta × 400
Fig. 64 hind foot × 400
Fig. 65 Abd VI anal spines and dorsal setae × 320
Figs. 66-70 Xenylla corticalis Boerner, drawn from specimen in Boerner collection at BMNH.
Fig. 66 hind foot × 400
Fig. 67 anal spine from side × 320
Fig. 68 SO Ant III × 1000
Fig. 69 apex of mucrodens, posterior view × 1000
Fig. 70 mucrodens × 320
Figs. 71-72 Xenylla grisea Axelson, drawn from a cotype in BMNH.
Fig. 71 hind foot × 400
Fig. 72 mucrodens × 320
Xenylla longicauda Folsom, 1898
I have examined specimens of this species in the collection of BMNH identified by Boerner as well as cotypes No. 5075 in USNM and from these two series the figures presented here were drawn. Folsom's original drawings lacked much of the fine detail shown here.
Notes: Claw and mucro finely granulated basally, Ant IV with apical sensory knob in pit, probably exsertile, and sensory rods and clubs as shown in Fig. 73. SO Ant III has two short sense clubs behind a deep cuticular fold which completely obscures them from the front or face of the organ, and one larger exposed sense rod on each side with two guard setae. Mucro varies from equal in length to about 0.75 length of dens and in no specimens is mucro larger than dens, as suggested by Folsom. The mucro (Figs. 77 and 80) has a small elevated lamella reaching about one-quarter down from base. Dens with two posterior setae only. Anal spines small on very small widely separated papillae, the spines (Fig. 82) subequal in length to 6-7 cuticular granules. The setae normally simple but the larger ones may be serrated.
Xenylla maritima Tullberg, 1869
These two figures drawn from a specimen in the BNMH identified by Boerner supplement those of Stach (1949 pl.xxiii).
Xenylla mucronata Axelson, 1903
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.page 18
Figs. 73-82 Xenylla longicauda Folsom, drawn from specimen in Boerner collection BMNH and from cotype in USNM.
Fig. 73 Ant IV and apex Ant III from cotype USNM showing sensory structures × 500
Fig. 74 hind foot from specimen in BMNH × 500
Fig. 75 ocelli from specimen in BMNH × 500
Fig. 76 anal spines from specimen in BMNH × 500
Fig. 77 mucrodens from specimen in BMNH × 500
Fig. 78 hind foot from cotype USNM × 600
Fig. 79 SO Ant III side view from cotype USNM × 1000
Fig. 80 mucrodens from cotype USNM × 600
Fig. 81 side view of mucro from cotype USNM × 600
Fig. 82 anal spines and associated setae from above cotype USNM × 1000
Figs. 83-84 Xenylaa maritima Tullberg, drawn from specimen in Boerner collection BMNH.
Fig. 83 hind foot × 400
Fig. 84 mucrodens × 400
Figs. 85-89 Xenylla mucronata Axelson, drawn from specimens in Boerner collection BMNH.
Fig. 85 apex Ant IV × 320
Fig. 86 hind foot × 400
Fig. 87 SO Ant III from side × 1000
Fig. 88 anal spines and dorsal setae on Abd VI × 320
Fig. 89 mucrodens × 1000
Xenylla rhodesiensis Womersley, 1926
When Womersley (1926, p.153) described this species the figures he included were rather inadequate. Specimens labelled cotypes are in the BMNH and I was able to prepare new figures from these.
Notes: The tenent hairs on the feet are 1,2,2, not 2,3,3, as given by Womersley; the large central seta of the tibiotarsus (Fig. 92) is not a clavate tenent hair. The sensory structures of Ants III and IV are as shown in Figs. 93 and 94. The anal spines (Fig. 91) are very small, not bigger than three cuticular granules. Rami of tenaculum each with three barbs. Length to 0.6 mm.
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).
Xenylla welchi Folsom, 1916
Specimens labelled cotypes No. 19904 in the USNM were used to prepare these drawings which supplement those given by Folsom with his original description.
Notes: Only the front feet have a single tenent hair, the others have two each. The mucro (Figs. 102-104) varies in outline between specimens. The minute anal spines are on papillae having very fine granules (Fig. 105). Ant IV with small exsertile knob, numerous short and long tapering sense rods but no stout bent sense rods.page 21
Figs. 90-95 Xenylla rhodesiensis Womersley, drawn from cotypes in BMNH.
Fig. 90 Mucrodens × 1000
Fig. 91 Abd VI with anal spines and dorsal setae × 400
Fig. 92 hind foot × 400
Fig. 93 SO Ant III × 1000
Fig. 94 apex Ant IV × 1000
Fig. 95 fore foot × 400
Figs. 96-100 Xenylla schillei Boerner, drawn from cotype in BMNH.
Fig. 96 anal spines and dorsal setae on Abd VI × 400
Fig. 97 apex Ant IV × 1000
Fig. 98 SO Ant III × 1000
Fig. 99 hind foot × 400
Fig. 100 mucrodens × 400
Figs. 101-105 Xenylla welchi Folsom, drawn from cotypes in USNM.
Fig. 101 fore foot × 1000
Fig. 102 posterior base of mucro × 1000
Fig. 103 mucrodens another specimen × 1000
Fig. 104 mucro another specimen × 1000
Fig. 105 anal spine and seta from side × 1000
Hypogastrura armata communis (Folsom, 1898) Nov. Comb.
Achorutes communis Folsom, 1898
Ceratophysella communis Stach, 1949
Ceratophysella yuasai Yosii, 1954
The specimen used to prepare these figures came from the Boerner collection BMNH and may be a cotype sent to Boerner by Folsom. C. yuasai Yosii comes from the same locality, Okayama in Japan, as Folsom's material and from Yosii's description and figures I conclude that C. yuasai Yosii is a subjective synonym of Folsom's species. In my "Index" in 1964 p.209 I placed A. communis Folsom as a subspecies of H. armata (Nicolet), and C. yuasai Yosii as a synonym of this subspecies H. armata communis (Folsom). Although I could not detect the sac-like organ between Ants III and IV on the specimen in the BMNH collections, Yosii in his description of C. yuasai states clearly that this organ is present. In accepting this I conclude that the species C. yuasai and A. communis are the same and, therefore, the nomenclatural changes set out above are correct.
Notes: Apex of Ant IV (Fig. 108) with sensory knob in a pit protected by an overlapping thickened cap and a dome-like sensory swelling. Sub-apically Ant IV has 6-7 long curved sense rods and numerous simple setae. Dens with 6-7 stout simple setae along posterior face.
Hypogastrura cavicola (Boerner, 1901)
The holotype of this species is in the Boerner collection BMNH and the accompanying figures were drawn from it. Stach (1949, p.110) gives further figures drawn from Polish specimens of this species.
Notes: Ant IV has an apical retractible papilla guarded by several slender short curved setae; 4-6 strongly bent, stout, exposed sense rods subapically and numerous simple setae. Anal spines on very stout stumpy contiguous papillae, the spines slightly longer than the hind claw. PAO with an associated "nebenhocker-like" structure (Fig. 115).page 23
Figs. 106-111 Hypogastrura armata communis (Folsom), drawn from specimen in Boerner collection in BMNH.
Fig. 106 mucro and apex of dens
Fig. 107 SO Ant III × 1000
Fig. 108 apex of Ant IV × 1000
Fig. 109 anal spines with associated setae × 400
Fig. 110 hind foot × 400
Fig. 111 PAO and adjacent ocelli × 1000
Figs. 112-115 Hypogastrura cavicola (Boerner) drawn from holotype in BMNH.
Fig. 112 hind foot × 400
Fig. 113 anal spines and setae from above × 400
Fig. 114 mucro × 1000
Fig. 115 PAO with associated "nebenhocker" and anterior ocelli × 1000
Hypogastrura gracilis (Folsom, 1899)
Achorutes gracilis Folsom, 1899
Achorutes tullbergi Schaeffer, 1900
Neogastrura aequepilosa Stach, 1949
Hypogastrura gracilis Yosii, 1960
Specimens of this species labelled cotypes are in the USNM and two specimens on slide No. 5078 were used for the preparation of these drawings.
Notes: SO Ant III as in Fig. 117. Ant IV subapically with trilobed sensory knob, 4 stout curved sense rods, numerous tapering, slightly curved, sense rods, and longer setae (Fig. 118). Front feet (Fig. 121) with two tenent hairs, others with three, all in line across tibiotarsus; base of claw and unguiculus granulated; PAO with four horizontal lobes and an erect lobe rising above the others (Fig. 116); rami of tenaculum with three barbs, corpus naked; dens (Fig. 119) posterior face coarsely granulated, the remainder much finer, the posterior face with five simple setae. Mucro (Fig. 120) with finely granulated broad inner lamella and small plain basal lamella; anal spines small, and separated on subequal papillae.
From my study of this material and of the descriptions given by Schaeffer (1900), Folsom (1899), Stach (1949) and Yosii (1960), I have concluded that both Schaeffer's and Stach's material is conspecific with that of Folsom's.
Hypogastrura humi (Folsom, 1916)
Achorutes humi Folsom, 1916
Specimens labelled cotypes in USNM on slide No. 14899 were used for the preparation of these drawings.
Notes: My drawing of the SO Ant III (Fig. 125) differs slightly from that of Folsom in the displacement of the sensory rods. The three tenent hairs of each tibiotarsus are arranged with the middle one arising above the others (Fig. 123); Ant IV with apical sensory knob, sense rods and setae arranged as in Fig. 124; dens with six simple setae on posterior face (Fig. 128); mucro with posterior face finely granulated and long simple inner lamella (Figs. 127 and 128).page 25
Figs. 116-122 Hypogastrura gracilis (Folsom), drawn from cotype in USNM.
Fig. 116 PAO and adjacent ocelli × 1000
Fig. 117 SO Ant III × 1000
Fig. 118 apex Ant IV × 1000
Fig. 119 mucrodens from side × 1000
Fig. 120 mucro and apex of dens of another cotype from side × 1500
Fig. 121 fore foot × 1000
Fig. 122 anal spine and papilla from side × 1000
Figs. 123-131 Hypogastrura humi (Folsom), drawn from cotypes in USNM.
Fig. 123 hind foot × 1000
Fig. 124 apex Ant IV × 1000
Fig. 125 SO Ant III × 1000
Fig. 126 tenaculum × 1000
Fig. 127 mucro and apex of dens from side × 1000
Fig. 128 mucrodens × 1000
Fig. 129 anal spine and associated seta from side × 1000
Fig. 130 PAO × 1000
Fig. 131 ocelli and PAO × 600
Hypogastrura macgillivrayi (Folsom, 1916)
Achorutes macgillivrayi Folsom, 1916
Specimens labelled cotypes on slide No. 19900 in USNM were used to prepare these drawings.
Notes: SO Ant III (Fig. 134) has sense clubs, sense rods and guard setae and is much more complicated than was indicated by Folsom; apex of Ant IV as in Fig. 135 with apical sensory knob, stout bent, and longer tapering, sense rods but only occasional setae; PAO variable with 4-5 peripheral lobes which may vary between opposite sides of one individual (Figs. 132 and 133); mucro with broad granulated lamella (Fig. 138); dens with six setae on posterior face (Fig. 137); two or three tenent hairs to each front foot (Fig. 136), three to each of the other feet, always in line across each tibiotarsus.
Hypogastrura myrmecophila Womersley, 1926
A specimen labelled cotype in BMNH was used to prepare these drawings which are additional to those given by Womersley (1926, p.156).
Hypogastrura nivicola (Fitch, 1847)
Podura nivicola Fitch, 1847
Achorutes socialis Uzel, 1890
Achorutes nivicola Folsom, 1902
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).page 28
Figs. 132-139 Hypogastrura macgillivrayi (Folsom), drawn from cotype in USNM.
Fig. 132 PAO and adjacent ocelli left side of animal × 1000
Fig. 133 PAO right side of animal × 1000
Fig. 134 SO Ant III × 1000
Fig. 135 apex Ant IV × 1000
Fig. 136 hind foot × 1000
Fig. 137 mucrodens × 1000
Fig. 138 mucro from another cotype × 1000
Fig. 130 anal spine and papilla from side × 1000
Figs. 140-143 Hypogastrura myrmecophila Womersley, drawn from cotype in BMNH.
Fig. 140 hind foot × 400
Fig. 141 ramus of tenaculum × 1000
Fig. 142 PAO and adjacent ocellus × 1000
Fig. 143 mucro from side × 1000
Hypogastrura pseudarmata (Folsom, 1916)
Achorutes pseudarmatus Folsom, 1916
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).page 30
Figs. 144-150 Hypogastrura nivicola (Fitch), drawn from cotypes in USNM and from a specimen in the Boerner collection BMNH.
Fig. 144 mucro and apex of dens × 1000 specimen BMNH
Fig. 145 mucrodens × 400 specimen BMNH
Fig. 146 hind foot × 400 specimen BMNH
Fig. 147 hind foot × 1000 cotype USNM
Fig. 148 PAO "Nebenhocker" and ocelli right side of animal × 600 cotype USNM
Fig. 149 PAO nebenhocker-like structure and adjacent ocelli × 1000, cotype USNM
Fig. 150 mucrodens × 1000, cotype USNM
Figs. 151-157 Hypogastrura pseudarmata (Folsom), drawn from co-types in USNM.
Fig. 151 Ant IV × 600
Fig. 152 PAO and adjacent ocelli × 1000
Fig. 153 PAO and ocelli right side of animal × 600
Fig. 154 mucrodens × 600
Fig. 155 hind foot × 1000
Fig. 156 SO Ant III × 1000
Fig. 157 Abd VI anal spines and setae from above × 600
Hypogastrura purpurascens galiciana (Boerner, 1903)
Achorutes purpurascens galiciana Boerner, 1903
The type and specimens labelled cotypes of this subspecies are preserved in the BMNH but no illustrations of it have previously been published.
Notes: Mucro (Fig. 160) is relatively longer and more tapering than in H. purpurascens s.str: it tapers apically to a point without the slightly recurved apex characteristic of H. purpurascens s.str; base of claw finely granulate; the unguiculus of the fore feet (Fig. 158) has the lamella considerably reduced as compared with the other feet (Fig. 159); the fore feet have two tenent hairs, the other feet three each, all in line across the tibiotarsus; rami of tenaculum with three barbs; the four segments of the antennae related as 20:23:27:45; SO Ant III normal with two sense clubs, two sense rods and three guard setae; Ant IV with apical sensory papilla, and 5-7 subapical stout curved sense rods; PAO with 3-4 peripheral lobes (Figs. 161-162): length of type specimen 2 mm.
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
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 page 33lamellate 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.page 34
Figs. 158-163 Hypogastrura purpurascens galiciana (Boerner), drawn from type and cotypes in BMNH.
Fig. 158 fore foot × 400 from type BMNH
Fig. 159 hind foot × 400 from type BMNH
Fig. 160 mucro apex of dens × 400 from type BMNH
Fig. 161 PAO × 1000 from type BMNH
Fig. 162 PAO and adjacent ocellus from cotype BMNH
Fig. 163 anal spine and setae from side × 400 from cotype BMNH
Figs. 164-171 Hypogastrura (Cyclograna) boletivorus (Packard), drawn from paratype of Cyclograna vulgaris.
Fig. 164 PAO surface appearance × 2000
Fig. 165 PAO deep appearance at base of organ × 2000
Fig. 166 SO Ant III × 2000
Fig. 167 apex Ant IV × 2000
Fig. 168 hind foot × 600
Fig. 169 mucrodens from side × 600
Fig. 170 mucrodens angled view × 600
Fig. 171 anal spine and associated setae × 600
Schaefferia variabilis Christiansen, 1951
The holotype and specimens labelled cotypes of this species are in USNM, and these drawings made from the type material will supplement those given in the original description, some of which are difficult to decipher from having been over-reduced in publication.
Proxenyllodes reticulatus (Boerner, 1909)
Hypogastrura reticulata Boerner, 1909
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
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.page 37
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
Figs. 183-189 Proxenyllodes reticulatus Boerner, drawn from the type in the Boerner collection at BMNH.
Fig. 183 Ant IV × 1000
Fig. 184 hind foot × 1000
Fig. 185 SO Ant III × 1000
Fig. 186 PAO and adjacent ocelli × 1000
Fig. 187 reticulated cuticle around the ocellar group and PAO, left hand side of animal × 400
Fig. 188 anal spines and papillae × 400
Fig. 189 mucro and apex of dens from side × 400
Figs. 190-198 Triacanthella alba Carpenter, drawn from paratypes in BMNH.
Fig. 190 PAO and ocellar group × 400
Fig. 191 PAO from other side of same specimen × 1000
Fig. 192 SO Ant III × 1000
Fig. 193 front foot × 400
Fig. 194 front foot from anterior face × 400
Fig. 195 anal spines and setae from side × 400
Fig. 196 anal spines and setae from another cotype specimen × 400
Fig. 197 mucrodens × 400
Fig. 198 mucro and apex dens × 1000
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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.