2001.J. Linn. SOC., 55: 1-52. With 33 figures
August 1974
The type specimens and identity of the species
described in the genus Lithobius by F. Meinert,
and now preserved in the Zoological Museum,
Copenhagen University
(Chilopoda: Lithobiomorpha)
E. H. EASON
Bourton Far Hill, Moreton-in-Marsh, Gloucestershire, GL56 9TN, England
Accepred for publication January I974
The 32 species described by Meinert in the genus Lithobius which are represented by type
material in t h e Zoological Museum, Copenhagen University, are redescribed from that material.
I t is concluded that 17 of these species are valid, one is a subspecies of Lithobius forficants (L.)
and 12 are junior synonyms: two are junior homonyms, but both are also synonyms of
described species.
CONTENTS
. . . . . .
Introduction
1. Lithobius intrepidus Meinert
2. L. borealis Meinert
. . .
3. L. microps Meinert
. . .
4. L. imperialis Meinert
. .
5. L. validus Meinert
. . .
6. L. voraxMeinert
. . .
7. L. romanusMeinert
. .
8. L. rhaeticus Meinert
.
9. L. rricuspisMeinert . . .
10. L. granulatus Meinert
.
11. L. obscurus Meinert
. .
12. L. rugosus Meinert
. . .
13. L. insignisMeinert
. . .
14. L. tenebromsMeinert
. .
1 5 . L. gracilipes Meinert
. .
16. L. puleher Miinert
. . .
17. L. bonensisMeinert
. .
18. L. grossidens Meinert
. ,
19. L. rapaxMeinert
. . .
20. L. suevicusMeinert . . .
21. L. hispanicus Meinert
.
22. L. lapidicola Meinert
.
23. L. eximius Meinert
. . .
24. L. audaxMeinen
. . .
25. L. flavusMeinert
. . .
26. L. gmcilis Meinert
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2
5
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23
25
26
26
27
27
30
31
34
36
37
38
40
E. H. EASON
2
2 7 . L. pyrenaicus Meinerr
28. I.. lutroMeinerr . .
29. 1.. latebrirola Meinert
3 0 . I.. galatheae Meinert
3 1. L. exiguus Meinert
.
32. L . IundiiMeinert
.
Acknowledgenients
. .
References
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. 41
42
. 43
. 44
. 45
. 47
48
48
INTRODUCTION
Of the 3 5 species described by Meinert in the genus Lithohius from 1868 t o
1886, three, namely L. latzelii, L. rantabrigensis and L. jowensis, are based on
specimens preserved in the Museum of Comparative Zoology at Harvard
College, Cambridge, Mass., U.S.A. (Meinert, 1886a). The type specimens of the
remaining 3 2 species are preserved in the Zoological Museum, Copenhagen
University: these have never been redescribed and in the present study an
attempt is made to establish their identity.
Although Meinert’s descriptions of these species are fairly adequate
according to the standards of his day, only about half of them have been
correctly identified by subsequent authors: of the remainder some have been
more or less disregarded while others have had their names misapplied leading
to a great deal of nomenclatural confusion. Of the 17 species shown here to be
valid, four were based on single specimens (holotypes) and 1 3 on more than
one specimen (syntypes). Lectotypes were selected from six of these syntypical
series by Miss D. Schmidt (Mrs Tobias) of the Senckenberg Museum,
Frankfurt-am-Maine, who examined most of Meinert’s material in 1960; and
from two of these series by Dr R. E. Crabill of the Smithsonian Institution,
Washington, who examined the material belonging t o the American species,
also in 1960: although none of this work was published it has been of great
assistance and in all cases Schmidt’s and Crabill’s selections are confirmed.
Lectotypes are also selected from four syntypical series which were examined
by neither Schmidt nor Crabill and a neotype is designated for L. borealis, the
type material of which has been destroyed.
Meinert’s practice (or the practice of whoever else may have arranged his
collection) was t o assemble together all the specimens, from whatever locality,
which he believed to belong to the same species and to place them in the same
tube under a composite locality label, so that the exact locality of some of the
lectotypes is unknown. Most of the Italian specimens were collected either
from Rasa (Razzes) in South Tirol or from the neighbourhood of Rome and
the adjacent Sabini Mountains: since South Tirol was part of Austria-Hungary
in Meinert’s day Rasa is not included among the Italian localities.
in this study full descriptions are given of specimens whose identity might be
controversial or which belong to species never adequately redescribed:
otherwise an account is given only of the more variable characters or those of
particular diagnostic importance, and reference is made to a previously
published description. Synonymies are given for the valid species only, and
these are restricted to synonyms incorporating a different specific name: those
incorporating no more than a change in the name of the genus or subgenus are
omitted, except in those few cases where the taxon in question has become
widely recognized. Conclusions as to the status and currently accepted generic
classification of all the species considered here are summarized in Table 1 .
SyntYPes
Neotype
Lectotype
Faeroe Is.
U.S.A.
1872
1868
1886
1872
1872
1886
1872
1872
1872
1872
1872
1872
1872
1868
1872
L. bonensis
L. borealis
L. exiguus
L. eximius
L. flavus
L. galatheae
L. gracilipes
L. gracilis
L. granulatus
L. grossidens
L. hispanicus
L. imperialis
L. insignis
L. intrepidus
L. lapidicola
inermir L. Koch, 1856.Syn. nov.
psacadonotus Attems, 1952.Syn nov.
Germany;
South Tirol; Italy;
Spain; Algeria
Denmark
Lectotype
Syntypes
Syntypes
Holotype
Italy
Spain
Lectotype
Spain
Syntypes
pilicornis
= L.
valid
pusillus pusillifrater Verhoeff, 1925.Syn. nov.
= L. macilentus L. Koch, 1862
= L. rubriceps
valid
= Polybothrus electrinus Verhoeff, 1934. Syn. nov.
= L. inops Brolemann, 1932.Syn. nov.
= L. mundanus Verhoeff, 1937.Syn. nov.
= L. alicatai Matic, 1967. Syn. nov.
valid
= L.
non L. granulatus L. Koch, 1862
= L. obscurus Meinert, 1872.Syn. nov.
Holotype
Uruguay
Nicobar Is.
= L. rubriceps Newport, 1845
non L. gracilis Porat, 1869
= L. guadarramus Matic, 1968.Syn. nov.
Holotype
Spain
= L. pilicornis Newport, 1844
= L.
= L.
= L . castaneus
Syntypes
Holotype
Madeira
S.S.
Lithobius
S.S.
Eupolybo thrus
(Parapolybothrus)
Lithobius
Paitobius
valid
= L. cantabrigensis zinus Chamberlin, 1911 ?
Lithobius S.S.
forficatus pulcher Meinert, 1872.Syn. nov.
Generic
classification
valid
= L. lapidicola: Latzel, 1880 (non Meinert, 1872)
= L. lusitanus wurmanus Verhoeff, 1937.Syn. nov.
= L. saalachiensis Verhoeff, 1937.Syn. nov.
= L.
= L . castaneus Newport, 1844
Validity and status
Spain
Holotypes
Spain
SYntYPe
Syntypes
South Tirol
Algeria
1872
L. audax
South Tirol; Italy;
Spain; Algeria;
Tunisia
Designate type
material
Country Of
localities
Date
Nominal species
Table 1
w
Syntypes
Lithobius S.S.
Bothropolys
Lithobius s.s
Lithobius S . S .
Lithobius S . S .
Neolithobius
valid
valid
= L . sydneyensis Pocock, 1891. Syn. nov.
subspecies of L. forficarus (Linn. 1758). Comb. nov.
valid
valid
= L . stejnegeri Bollman. 1893. Syn nov.
= L . tricuspis Meinert, 1872 (see p. 16)
valid
valid
= L . asperatus L. Koch, 1878, Syn. nov.
=I.. mutabilis L. Koch, 1862
valid
= L . nigrifrons Latzel & Haase, 1880. Syn. nov.
= L. falteronensis Manfredi, 1936. Syn. nov.
valid
valid
= L. brachycephalus Fanzago, 1880. Syn. nov.
valid
Holotype
Lectotype
Holotype
Lectotype
Lectotype
Syntypes
Lectotype
Lectotype
Syntypes
Holotype
Lectotype
Lectotype
Lectotype
Denmark
Spain
Algeria
Spain
Sakhalin (U.S.S.R.)
South Tirol
Italy
Hawaiian Is.
Germany
South Tirol
Germany;
South Tirol; Italy
South Tirol
U.S.A.
1868
1872
1872
1872
1872
1872
1872
1872
1872
1872
1872
1872
1872
L . microps
L. obscurus
L . pulcher
L. pyrenaicus
L . rapax
L . rhaeticus
L . romanus
L. rugosus
L. suevicus
L. tenebrosus
L. tricuspis
L . validus
L. vorax
Lithobius
(Ezembius)
Lithobius S.S.
Lithobius S.S.
Lithobius
(Monotarsobius)
valid
= Nampabius michiganensis Chamherlin, 1914. Syn. nov
U.S.A.
= L. duboscqui Brolemann, 1896
Lithobius S.S.
Nampabius
valid
Lectotype
Holotype
South Tirol
-
1872
Generic
classification
__ _ - - - __
1886
= I,. pyrenaicus Meinert, 1872. Syn. nov.
Validity and status
L . lundii
type
L. latro
-
Spain
-
___
1872
material
__ ___-L . late bricola
Country of type
localities
Date
Nominal species
Table 1 - a n t .
-
P
5
TYPE SPECIMENS O F LITHOBIUS
Note on Meinert’s labels
The exact wording on the original labels is recorded. The words following
the names of the species refer t o localities and are self-explanatory. These are
followed by the names of the collectors which, in the case of F. Meinert and V.
Bergsde, have been abbreviated to M. and Bgs. respectively. In addition to the
recorded words and letters most of the labels bear the letter “S” on the bottom
left-hand corner, probably indicating that the specimens in question were
placed in the collection by the entomologist Johann Schiddte (S. L. Tuxen,
in litt. ).
1. Lithobius intrepidus Meinert
Lithobius intrepidus Meinert, 1868: 262; 1872: 305.
Type localities. Maribo; Hellebaek; Mden (Denmark).
Type specimens. Syntypes: nine mutilated females, some immature, of L.
macilentus L. Koch, 1862 (= L. aulacopus Latzel, 1880) labelled “Lithobius
intrepidus Mein. Sjaell. Loll. MBen M.”
Description. In general agreement with Latzel’s ( 1880) and Brolemann’s
(1930) descriptions of L. aulacopus. The spinulation of the only specimen with
many legs intact is, however, in closer agreement with that of L. aulacopus
pyrenaicus Brolemann (Brolemann, 1930) than with that given by Brolemann for
the typical form of L. aulacopus.
Spin u la t ion :
Dorsal
Ventral
1
2
3
7
8
9
10-11
12
13
14
15
C
t
-
-
-
-
-
-
-
-
-
-
-
-
m
m
P
F
T
C
t
P
-
m
m
am
am
am
am
amp
amp
amp
amp
am
m
m
am
am
am
am
am
m
m
m
-
-
-
-
-
a
m
m
m
mp
amp
amp
amp
amp
-
-
-
-
-
-
-
-
F
a
aP
aP
P
aP
mP
aP
amp ap
amp aP
amp ap
m P P
m P P
m p p
T
a
aP
aP
aP
aP
ap
aP
ap
P
P
-
The 4th to 6th legs are all missing.
Remarks. Although synonymy of L. aulacopus Latzel with L. intrepidus was
recognized by Lohmander (1957), most authors have continued to use Latzel’s
name for this widespread European species. Eason (1972b) has shown that
both these names are junior synonyms of L. macilentus L. Koch.
2. Lithobius borealis Meinert
Lithobius borealis Meinert, 1868: 263 (non Meinert, 1872: 322).
6
E. H. EASON
Lithobius lapidicola: Latzel, 1880: 106 and most subsequent authors (non
Meinert, 1872: 328).
Lithobius erythrocephalus lapidicola: Verhoeff, 1925 : 146.
Lithobius lusitanus wunnanus Verhoeff, 1937: 212, 224, Fig. 54.
Lithobius saalachiensis Verhoeff, 1937: 214, 227, Fig. 51.
Non Lithobius borealis: Stuxberg, 1875a: 73; Verhoeff, 1937: 211.
Type locality. Faeroe Isles.
Type specimen. Neotype: owing to an accident for which the author was
responsible the syntypes of L. borealis have been destroyed: however, the type
locality of this species leaves no doubt as to its identity and a well-preserved
male, 10 mm long with 3 3 + 36 antenna1 articles and with the posterior angles
of ‘1.9 squared, has been selected from Faeroese specimens labelled
“Thorshavn, Uldspinderiet, under sten, 9.4.1925, Krygr” in the Zoological
Museum, Copenhagen University, and is here formally designated as the
neotype.
Description. Of the 136 mature examples of L. borealis from the Faeroe Isles
examined, all bear a fair resemblance to specimens of this species described
from Britain by Eason (1964) under L. lapidicola. Length: 9.0 to 10 mm.
Antennae: usually of 30 to 36 articles. Prosternum: usually with 2 + 2 teeth;
only two specimens, both females, have 2 + 3 teeth. Tergites: posterior angles
of T.9 squared, slightly projecting or, in a few specimens, quite markedly
projecting; those of T. 11 and 1 3 always with well-marked projections.
Spiizulation: presenting much the same range of variation as in British
specimens; supplementary prefemoral spines present on the 15th legs in 95%.
Further specimens. A male and a female in Meinert’s collection labelled
“Lithobius borealis Mein. 1872: belegmaterial, Cordoba, Bona M.” are clearly
the specimens on which he based his redescription of L. borealis (Meinert,
1872). They are not conspecific with the Faeroese specimens and both belong
to L. Eusitanus Verhoeff, 1925 (= L. borealis: Machado, 1952).
Remarks. The uncertainty surrounding the identity of this species has been
due to Meinert’s failure to note the supplementary prefemoral spine on the
15th leg in his original description and also to his redescription of L. borealis,
based entirely on examples of L. lusitanus from Spain and Algeria. Although in
Faeroese and northwest European specimens of L. borealis the posterior
projections on T. 11 and 1 3 are well-marked, in the central European form they
may be less obvious and this led Latzel ( 1 880) to describe Austro-Hungarian
examples, in which these projections are vestigial, under L. lapidicola Meinert
(see p. 34). At the same time Latzel was not altogether certain of the identity
of these specimens and, believing that they might indeed belong to L. borealis,
he sent some to Dr Meinert who identified them as L. lapidicola (Latzel, 1880:
108). I t thus seems that Meinert himself was guided more by the shape of the
tergites than by the characteristic spinulation and failed to identify Latzel’s
specimens with those he had originally described from the Faeroe Isles.
However, Latzel’s description is quite clear and mentions the supplementary
prefemoral spine, so that most subsequent authors have followed him in calling
the species L. lapidicola. Verhoeff (1925) discussed the confusion over the
identity of these two species, but added to it by making L. lapidicola (i.e.
borealis) a subspecies of L. erythrocephalus C. L. Koch. The same author later
TYPE SPECIMENS OF LITHOBIUS
7
(Verhoeff, 1937) used the name lapidicola for another species and described
the true L. borealis under L. saalachiensis (see also Jeekel, 1964a) and L.
lusitanus wiirmanus, differentiating these two forms by means of the shape of
the prosternum which is not an altogether reliable character.
Many of the accounts of L. borealis in the literature are not based on
examination of specimens but on copying one or other of Meinert’s
descriptions. Latzel (1880) and Verhoeff (1925) merely repeated Meinert’s
later (1872) description while Hammer (193 1) combined both the original
(1868) and the later. Brolemann (1930) based his account chiefly on Meinert’s
later description, supplementing it by the spinulation of a male from the north
of England: this specimen was fully described by Brade-Birks and Brade-Birks
(1933) and is probably an example of L. melanops Newport with the posterior
projections on T.9 imperfectly developed. Stuxberg’s (1875a) description of L.
borealis from Sweden was clearly based on examination of specimens but does
not seem to refer to the true borealis: although it agrees fairly well with
Meinert’s later description it is very unlikely that L. lusitanus occurs so far
north. Much the same can be said of Verhoeff‘s (1937) description of L.
borealis from Switzerland, Tirol and northern Europe. Lohmander ( 1 9 5 9 , the
only modern author to suggest the correct identity of L. borealis, believed that
both Stuxberg’s and Verhoeff‘s (1937) descriptions referred to a distinct
northern species to which he gave the name L. stella-martini. The same form
was described by von Porat (1889), but this author seems to have been entirely
dependant on Stuxberg for his account and the validity of L. stella-martini,
which has never been really adequately described, remains doubtful. Machado’s
(1952) description of L. borealis from Portugal was clearly based on examples
of L. lusitanus.
All that can be said with certainty is that most records of L. lapidicola in the
literature (see p. 34) refer to L. borealis, whereas most of those of L. borealis
refer either to L. lusitanus or to some other species.
In spite of having been originally described from the Faeroe Isles L. borealis
is widespread in central and southern Europe and is not essentially a northern
species. Scandinavian records are not numerous: Palmen (1 949) did not include
it in his list of Finnish species, Lohmander (1955) found it fairly widely but
only locally distributed in Sweden and it has been definitely recorded from
only two localities in southern Norway (B. Meidell, unpubl.).
3. Lithobius microps Meinert
Lithobius microps Meinert, 1868: 265; 1872: 3 3 0 (part).
Lithobius Duboscqui Brolemann, 1896: 116 and most subsequent authors.
Sigibius puntanus Chamberlin, 1913: 102.
Lithobius (Haplolithobius)olivarum Verhoeff, 1925: 152.
Monotarsobius duboscqui: Verhoeff, 1937: 189.
Non Lithobius (Monotarsobius) microps: Verhoeff, 1931 : 307 etc.
Type locality. Palm-house, Copenhagen Botanic Gardens (Denmark).
Type specimen. Holotype: shrivelled and mutilated immature female labelled
“Lithobius microps Mein. Kbhvn. bot. Have M.”
Description. Although this specimen is very defective there is no doubt that
8
E. H. EASON
it belongs to the species widely known as L. duboscqui, which has been fully
described by Brolemann (1930). Lertgth: 5.6 mm. Antemae: missing; with 3 3
articles according to Meinert. Ocelli: obscure but apparently two. Prosternum:
with 2 + 2 teeth. Tergites: without posterior projections. Coxal pores: 2,2,2,2.
15th leg: moderately swollen with single ventral median spines on the
trochanter, prefemur and femur, a single dorsal posterior spine on the
prefemur, and an accessory apical claw. Gonopod: with 2 + 2 unequal spurs
and a bidentate claw.
Further specimens. Nine very mutilated specimens in Meinert’s collection
labelled “Lithobius microps Mein. 1872: belegmaterial, Bona, Hispania M.”
seem to be those he used to supplement his redescription of L. microps
(Meinert, 1872). None of them are conspecific with the holotype, nor can they
be identified with any certainty.
Remarks. As in the case of L. borealis, the identity of L.microps has been
misunderstood owing t o Meinert’s redescription which was based on a number
of species from Spain and Algeria in addition to the true L. microps. In 1895
Brolemann recorded L. microps correctly from the Forest of Andaine in central
France; but in the following year, after studying Meinert’s redescription, ’le
Brolemann, 1896) decided that the specimens on which he had based this
record belonged to a new species, L. duboscqui. Thereafter duboscqui became
the almost universally accepted name for microps: only Lohmander (in Palmen,
1949), Machado (1952) and Jeekel (1964a) among modern authors have
recognized the identity of the two forms with one another. On the other hand
Verhoeff ( 1931, 1934, 1937 etc.) used the name microps for another species
(or group of species) which has never been fully described but which is
probably identical with (or probably includes) L. sukatus L. Koch from
Bavaria (Eason, 1972b): another species which has been overlooked by modern
authors but which may be identical with the microps of Verhoeff is L. biporus
Silvestri from the Italian peninsula. The species described by Brolemann (1930)
from the Maritime Alps under L. microps is clearly distinct from the true L.
microps, but there is some doubt as to its identity with the microps of
Verhoeff. Most other European authors have followed Verhoeff in their use of
this name so that records of L. microps Meinert in the recent literature are
largely misleading.
A further factor which has confused the identity of this species is the
variability in the number of antennal articles. Meinert originally recorded 3 3,
von Porat (1889) gave 25 t o 3 3 for specimens from Oslo (Norway), Palmen
(1949) gave 24 t o 29 (usually 25) for those from Finland, and Machado (1952)
gave 19 to 23 for those from Portugal. Brolemann (1896) recorded 23 t o 28
(usually 25) for L. duboscqui and later (Brolemann, 1930) described a
population from the Basses-Alpes with only 21 to 23. Jeekel (1964a) analysed
the number of these articles in specimens from three separate sites in northwest
Germany and found 22 to 24, 25, and 26 t o 29 respectively. Although most
British examples of this species have 23 to 27 (usually 25) articles (Eason,
1964), Bagnall (191 3) recorded two specimens from neighbouring localities in
the north of England with 32 and 25 antennal articles respectively: the first he
identified as L. microps and the second as L. duboscqui. But there is no
justification for making any taxonomic distinction between populations of this
species on the basis of the number of these articles. I t seems that those which
TYPE SPECIMENS O F LITHOBIUS
9
are exclusively synanthropic as in Scandinavia and much of northern Europe
contain a higher proportion of individuals with numerous antennal articles than
do those from further south.
The variability in the spinulation of the legs of L. microps has led to the
naming of a number of subspecies (of L. duboscqui) and one species, L.
olivarum Verhoeff from Italy, which Verhoeff placed in a new subgenus,
Haplolithobius; but few of these forms are worthy of taxonomic distinction.
Chamberlin ( 1 91 3) erected the genus Sigibius to receive a new species, S.
puritanus, based on specimens from the New England States (U.S.A.) which, as
Crabill & Lorenzo ( 1 957) suspected, belong to introduced but well-established
populations of L. microps (Crabill, personal communication). Sigibius, which
Chamberlin (1938) later regarded as a synonym of Haplolithobius and which
includes a number of species (Chamberlin, 195 2), differs from Monotarsobius
as originally described by Verhoeff (1905) in usually having 25 or more
antennal articles as compared with 19 to 22. Although Verhoeff’s (1937)
emended definition of Mono tarsobius would include Sigibius there seem to be
good grounds, quite apart from the number of antennal articles, for retaining
the distinction between the two taxa. However, the distinction has been
disregarded by European authors and Matic (1 970) made Sigibius a synonym of
Monotarsobius. Since it is not the purpose of this study to recommend changes
in established generic classification, L. microps is referred to Monotarsobius.
4.Lithobius imperialis Meinert
(Figs 1 to 3)
2
Figures 1 to 3. Eupolyborhius imperialis 1. Ocelli, right. 2. Ocelli, left. 3. Dental margin of
prosternum, right half, ventral.
Lithobius imperialis Meinert, 1872: 289.
Polybothrus (Parapolybothrus) electrinus Verhoeff, 1934: 65, Figs 74 & 75.
Eupolybothrus (Parapolybothrus) electrinus: Jeekel, 1967: 174.
Eupolybothrus (Parapolybothrus) imperialis: Jeekel, 1967: 174.
? Polybothrus vulcanius Verhoeff, 1942a: 492.
Type locality. Imperial ruins, Rome (Italy).
Type specimen. Holotype: a fairly well-preserved female labelled “Lithobius
imperialis Mein. Roma, Keiserborgen Bgs.”
Description. Size: 20 mm long and 2.8 mm broad at T.8. Colour: dull
yellow. Shape: breadths of head and of T.1, 3, 5, 8, 10, 12 and 14 to each
other as 64 : 64 : 60 : 65 : 70 : 69 : 66 : 54. Head: slightly broader than long;
E. H. EASON
10
lateral marginal interruptions distinct; posterior border barely emarginate,
almost straight. Antennae: 12.8 mm long; of 64 + 66 articles, transverse
proximally and elongate distally. Ocelli: 1 + 5, 4, 2, 3 (right) and 1 + 4 , 4 , 3, 2
(left) (Figs 1 and 2). Prosternuin: with 8 + 9 teeth and an incised median cleft;
Iateral spine very small, immediately posterolateral to lateral tooth (Fig. 3).
Tergites: moderately wrinkled and punctate; T. 1 trapeziform with posterior
border slightly emarginate; posterior borders of T.3, 5 , 8, 10, 12 and
intermediate tergite moderately emarginate, that of T. 14 strongly emarginate;
posterior angles of T.5 rounded, those of T.8 blunt, :hose of T.lO, 1 2 and 14
angulated but not sharp, those of T.4 squared, those of T.6 with small rounded
projections, those of T.7 with broad projections, those of T.9, 11 and 1 3 with
more prominent rather blunt projections. Coval pores: 20 to 30, mostly
circular and of very variable size, on each of the 12th to 15th coxae. 14th leg:
glandular pores concentrated on medial aspects of femur, tibia, tarsus and
metatarsus; without accessory apical claw. 15th leg: 14.4 mm long; glandular
pores as on 14th; without accessory apical claw. Clzaetotaxy of legs: setae
moderate in length and density on the first 1 3 pairs with the usual seriate
arrangement on the ventral aspects of tarsus and metatarsus; distal four articles
of 14th and 15th almost glabrous with a very few short setae. Gonopod: with
two subequal conical spurs separated from one another at their insertions by
their own breadth; claw simple.
Dorsal
Ventral
C
t
P
F
T
C
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
am
1
2-3
-
-
-
..
4
-
5-1 1
12-13
-
-
-
rn
14
a
a
m
15
rn
P
F
?
-
-
-
-
-
-
amp
amp
amp
amp
amp
amp
amp
a
ap
ap
ap
ap
a
a(p)
ap
ap
P
P
-
a
a
a
a
~
t
~
~
~
~~
a
p
~~
Vpl’ appears to b e a true spine and not an enlarged seta as in Eupolybothrus grossipes
(C. L Koch) (see Eason, 1970). 4 DpT is present on one side only.
Remarks. Jeekel ( 1967) referred L. imperialis tentatively to Parapolybothrus, a subgenus of Eupolybothms, but it has not hitherto been
redescribed: Fedrizzi’s (1877) brief account of the species is merely a
repetition of Meinert’s description. There is little doubt that the holotype of L.
imperialis is identical with Eupolybothrus (Parapolybothrus) electrinus
(Verhoeff), originally described from Teramo in the Italian peninsula, despite
the fact that some of the distinctive specific and subgeneric characters are
found only in males. There is nothing significant in Verhoeff‘s brief description
of Polybothrus vulcanius, based on a single female from Ischia, to separate it
from electrinus, but as Matic (1967) pointed out, Verhoeff contrasted vulcanius
with Eupolybothrus (s.s.) apenninigenus (Brolemann) which suggests that he
believed it to have quite different affinities, so that vukanius cannot be
proposed as a synonym of L. imperialis with complete confidence.
TYPE SPECIMENS OF LITHOBIUS
11
The form described by Manfredi ( 1956) as Polybothrus electrinus paulianus
from a cave in Campania (Italy) is merely a colour variant.
5. Lithobills validus Meinert
Lithobius validus Meinert, 1872: 291.
Lithobius riuizctulatus: Latzel, 1876: 97 and many subsequent authors.
Lithobius calabrensis Fanzago, 1880a: 269.
Lithobius brachycephalus Fanzago, 1880b: 16.
Lithobius Molleri Verhoeff, 1893: 3 17.
? Lithobius punctulutus C. L. Koch, 1847: 147.
Non Lithobius validus var. punctulatus: Latzel, 1888: 93; Verhoeff, 1900: 156
etc.
Type locality. Rasa (South Tirol).
Tilpe specimens. Lectotype: a well-preserved female selected by D. Schmidt
labelled “Lithobius validus Mein. Razzes M.” is here formally designated as the
lectotype. Paralecto type: a male accompanying the lectotype and conspecific
with it.
Description of lectotype. In general agreement with Latzel’s (1880)
description of L. validus and Prunescu’s (1966) description of L. punctulatus
punctulatus. Length: 25 mm. Head: sparsely and feebly punctate. Antennae: of
48 articles. Ocelli: 1 + 3, 4, 5, 5, 5 , 1. Prosternum: sparsely punctate; with
7 + 7 teeth of equal size and a small supplementary tooth adjacent to the
median cleft on one side; lateral spine small and pigmented. Tergites: T. 1 and 3
very sparsely and feebly punctate: posterior borders of large tergites and
intermediate tergite moderately emarginate; posterior angles of T. 5 rounded,
those of T.8 and 10 blunt, those of T.12 and 14 angulated, those of T . 6 , 7 , 9 ,
11 and 1 3 with projections. Coxal pores: 9, 8, 8, 7; oblong. Gonopod: as
figured by Prunescu (1966: P1. vi, C).
Spinulation:
Dorsal
Ventral
C
t
P
F
T
C
t
P
F
T
1
2-1 1
12-1 3
-
-
-
amp
amp
amp
amp
amp
a
ap
ap
P
p
a
ap
ap
P
15
am
am
am
am
am
-
-
amp
amp
amp
amp
amp
-
14
mp
mp
amp
amp
amp
-
-
m
m
m
a
a
a
-
-
-
-
Remarks. Although there has never been any doubt as to the identity of this
species there has been uncertainty as t o its correct name. Confusion arose when
Latzel (1876) described an example of L. validus under L. punctulatus and
later (Latzel, 1880), although using the name validus, stated that he believed
the species to be identical with L. punctulatus. Thereafter many authors
accepted L. validus Meinert as a junior synonym of L. punctulatus C. L. Koch,
but Eason (1972b) has suggested that L. punctulatus is more likely to refer to
Eupolybothrus grossipes (C. L. Koch) and that owing to the uncertainty
surrounding its identity the name punctulatus should be rejected as a nomen
dibium.
12
E. H. EASON
A further source of confusion was Latzel’s ( 1 888) use of the name L. validtrs
var. punctulatus C. Koch for a specimen from Bosnia lacking the posterior
projections on T.6. This led Verhoeff (1900, 1937) and some more recent
authors to use the name ptinctulatus for a southeast European form, probably
identical with Latzel’s Bosnian specimen, which they believed t o represent a
subspecies of validus. But Prunescu (1966) has shown that these southeast
European specimens belong to a distinct species, L. matici Prunescu, sympatric
with validus in the eastern part of its range. Some of the specimens from Bosnia
on which Verhoeff seems to have based his description of L. validus
ptrnctulatus are present in the Verhoeff Collection in the British Museum (N
H.) (Reg. no. 03.8.25.27-28): these have been examined and although Prunescu
was undoubtedly correct in regarding L. validus punctulatus (sensu Verhoeff)
as a distinct species, Verhoeff‘s specimens answer more closely to Prunescu’s
description of L. rnatici biharicus Prunescu than t o that of L. niatici matici.
Prunescu (1966) also reviewed a number of related forms from southwestern
and southeastern Europe which he believed to be conspecific with, though
subspecifically distinct from, L. punctulatus (i.e. validus). The “subspecies”
nzachadoi Prunescu from Portugal only differs from Meinert’s original
description of validus in having 50 to 5 3 antenna1 articles (Meinert gave 42 t o
44); but the lectotype of validus has 48 articles and up t o 5 1 have been found
in other examples of the species from the type locality (Rasa) preserved in the
British Museum (N. H.) (Reg. no. 13.6.18.614-615), so that machadoi should
be disregarded. The exact status of the other forms mentioned by Prunescu is
uncertain.
L. calabrensis from Italy was proposed as a synonym of L. validus by
Manfredi ( 1 957) and L. brach-vcephalza. also from Italy, was almost certainly
based on an aberrant example of L. validus. L. molleri from Portugal was
originally described as having three dorsal spines on the 15th femur and being
without puncta on either the prosternum or the tergites: the former feature is
so unusual in Lithobius that it can be dismissed as being due either to
aberration or to inaccurate observation, while the strength, density and
distribution of puncta in vulidus are much too variable to be of taxonomic
value. L. molleri should be regarded as a synonym of L. validits as proposed by
Brolemann (1930) and not as a separate species as Verhoeff (1937) continued
to maintain, nor as a subspecies of validus as proposed by Folkmanova (1947).
L. validits was the only species of Lithobiidae (as opposed to Ethopolidae)
included by Stuxberg (1875b) in the subgenus Eulithobius as he originally
defined it. Eulithobius has been used subsequently by both Chamberlin (1940)
and Crabill (1958) for a genus of North American Lithobiidae, but no type
species has been designated. Although, as Crabill pointed out, L. punctulatus
(i.e. validus) seems to be closely related t o these North American species, it is
also undoubtedly even more closely related to such European species as L.
variegatus Leach, L. rubriceps Newport and L. matici. Until the generic
classification of the European Lithobiidae has been revised, Etilitliobius should
be disregarded and L. validus should be referred to Lithobius S.S.
6. Lithobius vorax Meinert
Lithobius vorax Meinert, 1872: 292.
TYPE SPECIMENS OF LITHOBIUS
13
Neolithobius vorax: Chamberlin, 1925b: 480.
Type locality. Beloxi, near New Orleans (U.S.A.).
Type specimens. Lectotype: a rather mutilated male selected by R. E. Crabill
labelled “Lithobius vorax Mein. Beloxi, 1853 Kroyer” is here formally designed
as the lectotype. Paralectotype: a badly mutilated female accompanying the
lectotype and conspecific with it.
Description oflectotype. This specimen, 19 mm long, is in general agreement
with Chamberlin’s (1925b) detailed description of Neolithobius vorux. but it
has only 37 antenna1 articles (Chamberlin gave 38 to 43) and the prosternal
teeth, though essentially 5 + 5, are deformed on the left so that, counting from
the medial to the lateral, the second and fifth are bifid and the third trifid,
giving an apparent total of nine teeth.
Spinulation:
Ventral
Dorsal
C
t
P
F
T
C
14
a
a
15
a
m
m
m
m
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
amp
am
m
a
a
a
a
a
5-10
11
12
13
~
t
P
F
T
-
amp
amp
amp
amp
amp
amp
ap
ap
ap
p
P
p
ap
ap
ap
ap
P
-
-
-
~~
Owing to mutilation the spinulation of the first four pairs of legs could not be determined.
Remarks. Although Chamberlin’s (191la) early description of L. vorax
includes L. latzeli Meinert, he later (Chamberlin-, 1925b) showed these two
species to be distinct and designated L. vorax as the type species of
Neolithobius Stuxberg, a well-defined southern North American and Central
American genus.
7. Lithobius romanus Meinert
(Figs 4 and 5 )
Lithobius Romanus Meinert, 1872: 296.
Non Lithobius piceus romanus: Verhoeff, 1925: 138 etc.
Type localities. Imperial ruins and Campagna, Rome; Gennazano; Rocca di
Cavi (Italy).
Type specimens. Lectotype: a well-preserved male labelled “Lithobius
Romanus Mein. Italia Bgs.” is here formally designated as the lectotype.
Paralectotypes: eight males and three females, some immature, accompanying
the lectotype and conspecific with it.
Description of lectotype. General shape and appearance very similar to that
of L. piceus L Koch as described by Eason (1964). Length: 16.5 mm.
Antennae: of 40 articles, those of the distal half slightly elongate. Ocelli: 1 + 4,
3, 2; posterior ocellus little larger than posterosuperior. Prosternum: with 4 + 3
rather irregular teeth; lateral spine slender. Tergites: posterior borders of T. 1, 3 ,
5 , 8, 10 and 12 slightly emarginate, those of T.14 and intermediate tergite
E. H . EASON
14
0.5mm
0.1mm
1
Figures 4 and 5. Lithobius romanus 4. Right 0 gonopod, ventral. 5 . Left 9 genital claw, ventral.
more strongly emarginate; posterior angles of T.8 and 10 rounded, those of
T.12 blunt, those of T.14 angulated, those of T.9 and 11 with broad blunt
projections, those of T.13 with rather sharper projections; the shape of the
tergites is in marked contrast to that in L. piceus in which the posterior angles
of T.lO, 12 and 14 are sharp and slightly projecting; complete absence of setae
on the posterior border of T.9 distinguishes L. romanus from the closely
related L. capreae Verhoeff (see also Matic, 1971). Coxal pores: 4, 5, 5, 4;
circular. 14th and 13th legs: very slightly swollen, but no more so than in
females; transition in breadth of 15th leg at the tibio-tarsal articulation a little
less marked than in L. piceus; 15th accessory apical claw two-fifths the length
of principal claw. Genitalia: gonopod of a single article; second genital sternite
without setae.
Spinulation:
Ventral
C
1
2 -3
4-5
6
7 -9
10-12
13
14
15
t
P
Dorsal
F
T
C
t
am
am
am
am
(a)
a
a
m
m
m
am
am
-
-
am
am
a
a
a
a
-
13 VaC and 6 VaP are present o n one side only.
P
F
T
TYPE SPECIMENS OF LITHOBIUS
15
Description o f adult paralectotypes. Differing from the lectotype in the
following characters: Length: 17 to 20 mm. Antennae: of 32 to 42 articles.
Ocelli: 1 + 4, 3 , 2 to 1 + 4, 4, 3 , 2,. Prosternum: with 3 + 4 or 4 + 4 teeth.
Coxal pores: 4, 5, 5, 4; 5 , 6, 6 , 4; 6, 6, 6, 5 or 5 , 7, 7, 5 ; oval in larger
specimens. Female gonopod: with two unequal spurs; in one specimen the claw
is bidentate with the medial denticle well-developed (Fig. 4), in another the
claw is almost simple (Fig. 5). Spinulation: 14 VaC and 14 VmT may be
absent; 1 5 VmT, 1 DaP, 14 DaF and 1 DpT may be present; VaP may be
present on all the legs, may be discontinuous as in the lectotype, or may be
absent on the first nine legs.
Remarks. Although Meinert recorded this species from a number of localities
in the neighbourhood of Rome, it seems to have been overlooked by most
subsequent authors. Manfredi made no mention of it in any of her numerous
works on the Italian myriapod fauna and may have misdetermined it as L.
peregrinus Latzel for which it is easily mistaken. Verhoeff described, under L.
piceus romanus Meinert, a form of L. piceus L. Koch from Italy (Verhoeff,
1925) and Switzerland (Verhoeff, 1935) with three spurs and a simple claw on
the female gonopod. Demange (1958a) pointed out that this description does
not refer to L. romanus Meinert which has only two such spurs, but to a form
he had himself found in Switzerland which he described as a new subspecies, L.
piceus verhoeffi Some of the Italian specimens on which Verhoeff probably
based his description of L. piceus romanus are present in the Verhoeff
Collection in the British Museum (N. H.) (Reg. no. 1921.6.10.331-336): these
have been examined and their identity with L. piceus verhoeffi Demange is
confirmed. The identity of L. romanus remained in doubt until Matic (Matic &
Darabantu, 1968) redescribed it briefly but correctly and restored it to specific
status.
The distribution of L. romanus outside Italy is uncertain. A closely related
species, L. nigripalpis L. Koch (= bulgaricus Verhoeff) which is widely
distributed in the Balkans, the Aegean Archipelago and Asia Minor (Eason,
1972b), can only be distingushed with certainty from L. romanus by the more
slender spurs and sharper simple claw on the female gonopod. Attems’s (1902)
description of L. romanus from Crete, though agreeing with romanus in all
respects, does not include a detailed description of the female gonopod and
might possibly refer to L. nigripalpis. The same author’s description of L.
romanus from several localities in Syria (Attems, 1926) and his records of the
species from Angora and Smyrna in Turkey (Attems, 1932) almost certainly
refer to nigripalpis. If L. romanus is, in fact, confined to Italy it may be no
more than a subspecies of L. nigripalpis L. Koch, 1867.
8. Lithobius rhaeticus Meinert
Lithobius Rhaeticus Meinert, 1872: 297.
Type locality. Rasa (South Tirol).
Type specimens. Syntypes: two females and one male of L. tricuspis
Meinert, 1872 labelled “Lithobius Rhaeticus Mein. Razzes M.”
Description. Apart from presenting only two spurs
on each female gonopod
E. H. EASON
16
these specimens agree with Latzel’s (1880) and Brolemann’s (1930) descriptions of L. tricuspis. The female genital claw has a well-developed medial
denticle and a much smaller lateral denticle. The claw is therefore tridentate,
but in one of the females the lateral denticle is very inconspicuous and this, no
doubt, led Meinert t o describe the claw as bilobed.
Remurks. Fedrizzi (1878) recognized L. rhaeticus as a synonym of L.
tricuspis and Latzel (1880), while giving the same synonymy, suggested that
rhueticus might be retained as the name of a variety of L. tricuspis lacking the
small medial spur on the female gonopod. But the name rhaeticus has never
been used in this way by subsequent authors although the form with only two
female genital spurs on each side has been redescribed as L. tricuspis
dolomiticits Attems (Attems, 1927) and L. tricuspis strusseri Verhoeff
(Verhoeff, 1937), in both cases from South Tirol. Being sympatric with the
typical L. tricuspis this form cannot be a subspecies, but if it is thought
desirable t o give it varietal status the name rhaeticus should be used.
9. Lithobius tricuspis Meinert
LitFiobius RIiaeticus Meinert, 1872: 297.
Litlzobiits tricuspis Meinert, 1872: 298.
Litiiobius piceus tricuspis: Brolemann, 1898: 200.
Lithobius dolomiticus Attems, 1903: 113.
Type localities. Tubingen, Wurtemberg (Germany). Rasa (South Tirol).
Roman Campagna; Gennazano; Ariccia (Italy).
Type specimens. Lectotype: a well-preserved female selected by D. Schmidt
labelled “Lithobius tricuspis Mein. Tirol M. Italia Bgs.” is here formally
designated as the lectotype. Paralectotypes: nine females, eight males and an
agenitalis accompanying the lectotype and conspecific with it, together with a
male of L. mutabilis L. Koch, a male of L. nicoeensis Brolemann, three males
of L. aostanus Verhoeff?, an immature male of L. romanus ? and five immature
specimens which could not be identified.
Description of lectotype. In general agreement with Latzel’s (1880),
Brolemann’s (1930) and Eason’s (1965) descriptions of L. tricuspis. Length:
13.5 mm. Antennae: of 48 articles. Tergites: as described and figured by Eason
(1965: Fig. 3). Coxal pores: 3, 4,4,4 and 4, 4, 4, 4; circular. Gonopod: with
three somewhat spinous spurs as figured by Eason (1965: Fig. 5); claw with a
well-developed medial denticle and a much smaller, more proximal, lateral
denticle.
Spin ula tion :
Ventral
C
1
2
3
4-7
8-11
12
13-14
15
t
P
Dorsal
F
T
m
m
m
am
am
am
am
a
C
t
P
F
T
TYPE SPECIMENS O F LITHOBIUS
17
Gonopods o f female paralectotypes. The spurs are usually as in the lectotype
but in one specimen the lateral spur of both gonopods is feebly clubbed at its
extremity as in the German specimen figured by Verhoeff (193 5: Fig. 15) and
referred by him t o a new subspecies, L tricuspis belchenins: in another
specimen one of the gonopods lacks the small medial spur. The claw is also
usually as in the lectotype but in one specimen it is more frankly tridentate
with a well-developed lateral denticle, and in another it is bidentate without
trace of the lateral denticle as figured by Eason (1965: Fig. 5).
Further specimens. A female of L. Iwitanus crissolensis Verhoeff and a
female of L. upruzianus Verhoeff, together labelled “Lithobius tricuspis
monstr. Mein. Gennazano Bgs.” were clearly not intended by Meinert t o form
part of the syntypical series.
Remarks. Meinert’s inclusion of examples of so many different species
among the type specimens of L. tricuspis does not suggest that he examined
them all very carefully. However, the identity of the lectotype with the
well-known and widespread species known to all subsequent authors as L.
tricuspis is not in doubt. Although published on a later page of Meinert’s paper
than that of L. rhaeticus (see p. 15), his description of L. tricuspis was
considered by Latzel (1880), who was the first author to revise Meinert’s
European species, to be the more definitive: the name tricuspis is therefore,
regarded as having priority over rhaeticus (Code, article 24a).
Attems (1903) based L. dolomiticus on a single male from South Tirol: he
later (Attems, 1927) described a female and made dolomiticus a subspecies of
L. tricuspis; but this form, like rhaeticus, only differs from the typical L.
tricuspis in lacking the small medial spur on the female gonopod. In one of his
earliest works Brolemann ( 1898) made L. tricuspis a subspecies of L. piceus L.
Koch but this combination was not justified and has been disregarded by
subsequent authors including Brolemann himself.
The nominal subspecies and varieties of L. tricuspis have been reviewed by
Eason (1965). Few of these are valid although L. tricuspis var. mononyx Latzel
and L. tricuspis var. multidens Demange have been shown by Tobias (1969)
and Matic (Matic et al., 1967) respectively to belong to distinct species.
10. Lithobius granulatus Meinert
Lithobius granulatus Meinert, 1872: 299.
Non Lithobius granulatus L. Koch, 1862: 67.
Type locality. Montevideo (Uruguay).
Type specimen. Holotype: a fairly well-preserved male of L. obscztrus
Meinert, 1872 (= L. sydneyensis Pocock, 1891) labelled “Lithobius granulatus
Mein. Montevideo 618.47 Rk.H.”
Description. In general agreement with Brolemann’s (1924a) description of
L. aruichensis Brolemann & Eason’s (197 3 ) description of L. sydneyensis
Pocock. Length: 14 mm. Antennae: of 26 articles. Coxal pores: 5,6,6,4: oval.
15th leg: with the characteristic wart-like outgrowth on the femur as figured by
Eason (1973a: Fig. 35).
2
E. H. EASON
18
Spin ula t io ti.
Ventral
C
1
24
5 -9
10-12
13
14
15
t
P
F
T
C
t
F
T
am
am
am
am
am
a
a
In addition to the above there is a supplementary posterior prefernoral spine o n the left
14th leg and three such spines o n the right 15th leg: the left 15th leg is missing.
Rerizurks. Silvestri (1899) described some specimens from Santiago (Chile)
under L. platensis Gervais and later (Silvestri, 1905) recorded this same form
from Montevideo. Whether he was justified in using Gervais’s name is
questionable because L. platensis, although from Montevideo, gas originally
described as having very long antennae of 36 to 40 articles (Walckenaer &
Gervais, 1847) whereas Silvestri gave 24 to 26 articles. However, apart from his
failure to record the small accessory apical claw on the 15th leg which is easily
overlooked, Silvestri’s description agrees in all essentials with the holotype of
L. granulatus and had Meinert not overlooked the wart-like outgrowth on the
15th femur the identity of these two forms with one another would have been
recognized from their respective descriptions. On the other hand the holotype
of L. granulatus undoubtedly belongs to the western Mediterranean species
known to European workers as L. araichensis Brolemann which Eason (1973)
has shown to be a junior synonym of L. sydneyensis Pocock from Australia.
This latter name must now fall as a synonym to L. ohscurzis Meinert, the next
species to be considered, whose identity has not hitherto been suspected (see
below).
The occurrence of this species in South America has already been foreseen
(Eason, 1973) so that the identity of L. granulatus is not unexpected. The
name granulatus must be rejected as a junior homonym of L. granulatus L.
Koch, 1862.
11. Lithobius obsciirus Meinert
Lithobius granularus Meinert, 1872 : 299 (non Koch, 1862).
Lithobius obscttrus Meinert, 1872: 300.
Lithobius sydneyerisis Pocock, 1891b: 153.
Lithobius bermudensis Pocock, 1893: 126.
Lithobius platensis: Silvestri, 1899: 142.
Lithobius arakhensis Brolemann, 1924a : 58, Figs. 4-6.
Walesobius excrescens Attems, 1928: 78.
Chilebius coquimbo Chamberlin, 1955: 56.
? Lithobius platensis Gervais in Walckenaer & Gervais, 1847: 237.
? Kesubius syntheticus Chamberlin, 1925c: 120.
? Andebius calla0 Chamberlin, 1955: 57.
19
TYPE SPECIMENS O F LITHOBIUS
Type localities. Near Granada; Caratraca near Malaga (Spain).
Type specimens. Lectotype: a well-preserved female selected by D. Schmidt
labelled “Lithobius obscurus Mein. Hispan. mer. M., L.Lund” is here formally
designated as the lectotype. Paralectotype a rather mutilated female accompanying the lectotype and conspecific with it.
Description of lectotype. In general agreement with Brolemann’s description
of L. arakhensis and Eason’s (1973) description of L. sydneyensis. Length:
13.2 mm. Antennae: 5.2 mm long; of 26 rather elongate articles. Ocelli: 1 + 3,
4,2. Prosternum: as figured by Eason (1973: Fig. 34). Tergites: as figured by
Brolemann (1924a: Fig. 4). Coxal pores: 4, 5, 5 , 4;circular. 14th leg: very
slightly swollen; accessory apical claw well-developed. 15th leg: 5.2 mm long;
very slightly swollen; accessory apical claw very small. Gonopod: as figured by
Eason (1973: Fig. 36) but the spurs more closely set, separated from one
another at their insertions by barely their own breadth.
Spin ula ti0 n :
Dorsal
Ventral
1-2
3-10
11
12
13
14
15
C
t
P
F
T
C
t
P
F
T
-
-
am
am
am
am
am
a
a
-
-
-
a
aP
ap
ap
a
a
a
-
mp
mP
amp
amp
amp
amp
amp
ap
aP
ap
ap
m
m
m
amp
amp
amp
amp
amp
amp
am
-
-
mp
mp
mp
amp
amp
amp
amp
P
P
p
P
P
-
-
-
-
In addition to the above there are two supplementary posterior prefemoral spines on the
right 15th leg and one on the left 15th leg.
Remarks. Those authors who have included L. obscurus in their keys to
Iberian species (Brolemann, 1924b; Demange, 1958b; Matic et al., 1967) have
added nothing t o Meinert’s description and its identity has been overlooked
owing to Meinert’s failure to note the supplementary prefermoral spines on the
15th legs, and because the really characteristic feature of the species, the
wart-like outgrowth on the 15th femur, is only present in males.
The synonymy and distribution of this species have been fully discussed
under L. sydneyensis by Eason (1973) who suggested L. platensis (sensu
Silvestri) and Chilebius coquimbo, both based on males and almost certainly
identical with one another, and both from coastal localities in South America,
as possible synonyms. With the discovery of the identity of L. granulatus
Meinert from Montevideo (see p. 17) these two names can now be added to
the list of synonyms with confidence: the likelihood of these forms belonging
t o an indigenous South American species or to some introduced species other
than L. obscurus is altogether remote. The identity of the other South
American species tentatively suggested (Eason, 1973) as a synonym, Andebius
callao which was based on a female from Peru, is less certain. But if, as seems
likely, the indigenous species of Lithobiidae in South America are confined t o
the north of the continent (Eason, 1974) there is yet another species,
Kesubius syntheticus, which is a probable synonym. K. syntheticus was based
on a male from Uruguay with the 15th legs missing and a wart-like outgrowth
on the 14th femur. Males of Lithobius calcaratus C. L. Koch, a common
20
E. H. EASON
European species with comparable secondary sexual modification of the 15th
femur, are sometimes found with this same modification on the 14th femur,
but only when the corresponding 15th leg has been lost (Eason, 1972,b). I t
may be that loss of the 15th leg in certain circumstances induces, in subsequent
stadia, not regeneration of the leg but growth of the secondary sexual character
on the 14th leg: this phenomenon occurring in L. obscurus would result in an
individual closely resembling K . syrztheticzrs. The doubt as to the identity of
L. platensis Gervais has already been discussed (see p. 19).
The numerous genera other than Lithobitis which are represented among the
established and questionable synonyms of L . obsczirzw were all erected by
Chamberlin and are evidence of this author’s tendency to name genera on the
basis of very trivial characters (see also Eason, 1974).
12. Lithobius rugosus Meinert
(Figs 6 and 7)
I
7
L
0.5mm
I
Figures 6 and 7. Borhropdys rugoms 6. Ocelli. 7. Dental margin of prosternurn, right half,
ventral.
Lithobius rugosus Meinert, 1872: 306.
Lithobius asperatus L Roch, 1878: 788.
Lithobius Thetidis Karsch, 1880:848.
Lithobius shimensis Pocock, 1895: 349.
Bothropolys asperatus: Attems, 1914:
Bothropolys rugosus: Attems, 1914:
Bothropolys spinosior Chamberlin, 1920: 78.
Ethopolys rugosus: Chamberlin, 1920: 78.
? Bothropolys migrans Chamberlin, 1930: 69.
Type locality. Oahu, Hawaiian Islands.
Type specimens. Lectotype: a male with both 14th and 15th legs missing but
otherwise well-preserved labelled “Lithobius rugosus Mein. Oahu 10/96
Galatea” is here formally designated as the lectotype. Paralectotypes: four
adult males and one immature male, each with both 14th and 15th legs missing,
accompanying the lectotype and conspecific with it.
Detached 14th and 15th legs, probably belonging to the lectotype, have now
been placed in a separate tube.
Description of lectotype. Size: 21 mm long and 2.75 mm broad at T.lO.
TYPE SPECIMENS OF LITHOBIUS
21
Colour: dark brown. Shape: breadths of head and of T . l , 3 , 5 , 8 , 10, 12 and 14
to each other as 64: 66: 62: 64: 68: 69: 66: 54. Head: slightly longer than
broad; no lateral marginal interruptions; posterior border barely emarginate,
almost straight, with marked localized median thickening of marginal ridge.
Antennae: 9.6 mm long: of 20 elongate articles. Ocelli: 1 + 6, 6 , 4 , 3 , (Fig. 6).
Prosternum: with 7 + 7 teeth, the line of their apices markedly recurved, and a
narrow median cleft; lateral spine very small, immediately posterolateral to
lateral tooth (Fig. 7). Tergites: strongly wrinkled and coarsely punctate,
particularly T.8, 10, 12, and 14; T.l trapeziform with posterior border barely
emarginate, almost straight; posterior border of T.3 very slightly emarginate,
those of T.5, 8, 10, 12, 14 and intermediate tergite moderately emarginate;
posterior angles of T.5 and 8 rounded, those of T.10 and 12 blunt, those of
T.14 angulated, those of T.4 squared, those of T.6 with very feeble rounded
prqjections, those of T.7 with broad but not prominent projections, those of
T.9, 1 1 and 1 3 with very prominent narrow projections. Coxal pores: 23, 28,
26, 1 3 ; irregulary arranged in about three rows. 14th leg: not obviously
swollen; glandular pores concentrated on medial aspects of femur, tibia, tarsus
and metatarsus; femur with a feeble narrow dorsal sulcus throughout its length;
without accessory apical claw. 25th leg: 10 mm long; not obviously swollen;
glandular pores as on 14th; prefemur with a very feeble narrow dorsal sulcus;
femur with a more distinct dorsal sulcus on its distal half, narrow and shallow
proximally, deeper and broader distally; without accessory apical claw.
Chaetotaxy of legs: setae rather sparse on the first 1 3 pairs except for the
seriate setae on the ventral aspects of tarsus and metatarsus which are more
numerous; 14th and 15th almost glabrous with a few very short setae.
Genitalia: gonopod of a single article; second genital sternite without setae.
Spinula tion :
Dorsal
Ventral
1-2
3
4-5
6-10
11
12-13
14
15
T
C
t
P
F
-
-
-
-
mp
mp
(ahp
amp
amp
amp
amp
amp
amp am
amp am
amp am
amp am
amp am
amp am
a
am
a
am
am
am
-
m
m
m
C
t
P
a
a
a
a
-
amp
amp
amp
amp
amp
amp
amp
amp
-
-
F
T
4 and 5 VaP are present on one side only.
The spinulation and other characters of the 14th and 15th legs are taken
from detached legs which may possibly belong to the paralectotypes.
Description of adult paralectotypes. From 18 to 21 mm long and agreeing in
all essential characters with the lectotype, except that the head is usually as
broad as T. 1, and in one specimen the posterior projections on T.6 are vestigial.
Remarks. Stuxberg (1875b) proposed L. rugosus as a junior synonym of
Ethopolys xanti (Wood), a Californian species, but Chamberlin (1920),
although mistakenly referring rugosus to Ethopolys (no doubt because of
Meinert’s failure to note the posterior projections o n T.6 and 7), questioned
this synonymy. Haase (1887) gave a brief account of the species which is
22
E. H. EASON
merely a repetition of Meinert’s description. Attems (19 14) referred ricgosus
correctly to Rothropolys but he described the 15th coxa as being without the
lateral spine (VaC), probably owing t o misinterpretation of Meinert’s description which mentions only two 15th coxal spines. But Meinert’ disregarded the
dorsal spinulation of the legs in all his descriptions of Lithobiris species and the
two coxal spines he mentioned are VaC and VmC, and not DaC and VmC as
Attems supposed. I t seems that all these authors were dependent solely on
Meinert’s description of L rirgosirs for their accounts of this species and its true
identity has been overlooked.
The type specimens of L. rirgosirs all belong to the widely distributed eastern
Asiatic species known as Rothropol.vs asperatiis. This species, originally
described from Japan (L. Yoch, 1878), has been redescribed by Attems (1909)
and there is no doubt as to the identity of Meinert’s specimens. L thetidis from
China was proposed as a synonym of L asperatiis by Haase (1887), L.
shimensis from Japan by Eason (1973), and Bothropolys spinosior from Japan
also by Eason (1972b): all these names now become junior synonyms of L.
rugo sii 5
I t is unlikely that this species is indigenous in Hawaii and it has almost
certainlv been introduced either from Japan or the mainland of eastern Asia. A
closely similar species, Bothropolys niigraiis which may well prove to be
another synonym of nrgosw, was recorded from Hawaii as an immigrant,
introduced with plants imported from China (Chamberlin, 1930). The only
other species of Rothropolys from Hawaii, B rnaltihianics Attems, 1914 (= B
oahicanr~s Chamberlin, 1920), was originally recorded from Oahu as L.
uJperatzts by Attems (1903) but, assuming Attems’s description to be accurate,
it cannot be regarded as identical with B rtrgoszis owing to its very deficient
spinulation.
As well as occurring in eastern Asia the genus Bothropolys has an extensive
distribution in North America where the most widespread species, R.
rntrlticientatiis (Newport), is so close to R rirgostrs as to suggest the possibility
of the same species occurring in both continents. Rut Chamberlin’s (1925a)
detailed description of B rnrrltideritatirs shows that the two forms can be
distinguished by the shape and chaetotaxy of the male 15th legs as well as by a
few other less striking characters, and probably belong to separate species.
13. Lithobiirs irisigriis Meinert
Lithobills insignis Meinert, 1872: 3 1 3
T,vpe localitits. Escorial; Caratraca near Malaga (Spain).
Type specimens. Sjxtypes: three females, a male and an agenitalis of L.
rtrbriceps Newport, 1845 labelled “Lithobiirs insignis Mein. Hispania Collin, L.
Lund.”
Oescription. The adults, 22 to 26 mm long with 48 to 52 antenna1 articles,
15 to 20 ocelli, 6 + 6 or 7 + 7 prosternal teeth and 6, 5, 5, 5 circular, oval or
oblong coxal pores all agree with Machado’s (1952) description of L. insignis.
One of the females has distinct broad posterior projections on T.7 in addition
to the much more marked projections on T.9, 11 and 13. The degree of
punctation of the integument is variable: one specimen is quite densely
TYPE SPECIMENS OF LITHOBIUS
23
punctate on the head, prosternum and tergites as far back as T.12, whereas in
another the puncta are sparse and confined t o the head and prosternum.
Remarks. Those specimens with distinct puncta on the more posterior
tergites answer to L. insignis hessei Verhoeff (Verhoeff, 1928), and the female
with posterior projections on T.7 to L. insignis var. anomaluns Machado
(Machado, 1952). But these two features are much too variable to be of
taxonomic value and both hessei and aizomalans should be disregarded.
Eason (1972a) has shown that the correct name for this well-known Iberian
species is L. rubriceps Newport.
14. Lithobius tenebrosus Meinert
(Figs 8 to 11)
0.2 mm
Q5mm
I
Figures 8 to 11. Lirhobius fenebrosus 8. Ocelli. 9 . Dental margin of prosternum, ventral. 10.
Left 15th leg of 9, medial (posterior). 11. 9 gonopods, ventral.
E. H . EASON
24
Lithobius tenebrosus Meinert, 1872: 3 17.
Lithobius nigrifrons Latzel & Haase in Latzel, 1880: 71.
Litlzobius falteronensis Manfredi, 1936: 125.
T.vpe locality. Rasa (South Tirol).
Type speciinerz. Holotype: a fairly well-preserved female labelled “Lithobius
tei?ehrostu Mein. Razzes M.”
Description. Size: 11.8 mm long and 1.7 mm broad at T.lO. Colour: brown.
Shape. breadths of head and of T . l , 3 , 5, 8, 10, 12 and 14 to each other as
72 : 61 : 67 : 73 : 82 : 84 : 78 : 65. Head: broader than long; lateral marginal
interruptions distinct; posterior border very feebly emarginate. Antennae:
5.0 mm long; of 39 articles. Ocelli: 1 + 4, 4, 4 , 4 , 3 (Fig. 8). Prosternum: with
2 + 2 very prominent teeth, the medial pair the larger and the line of their
apices straight; lateral spine setiform (Fig. 9). Tergites: posterior borders of
large tergites and intermediate tergite only slightly emarginate; posterior angles
of T.5 and 8 rounded, those of T.10 and 12 blunt, those of T.14 angulated,
those of T.9, 11 and 1 3 with broad blunt projections. Coxal pores: 5 , 6 , 6, 5 ;
large and circular or somewhat oval; separated from one another by less than
their own diameter. 15th leg: 4.0 mm long; femur slightly swollen and feebly
bowed dorsally; without accessory apical claw (Fig. 10). Gonopod: with stout
conical spurs, two on the right gonopod and three on the left; claw with medial
and lateral denticles distinct; a line of four stout dorsolateral setae on the
second article (Fig. 11).
Spin ula tiori:
Ventral
C
1
2-3
4
5 -6
7 -9
10
11-12
13
14
15
t
P
Dorsal
F
T
c
t
am
am
am
am
am
am
amp
amp
amp
am
m
-
m
am
am
am
am
am
am
am
-
-
-
-
-
-
-
_.
-
-
a
a
a
-
-
P
F
T
P
mP
mP
mP
amp
amp
amp
amp
amp
amp
The median ventral spines tend to be longer than in most species of Lithobius (Fig. 1 0 ) .
1 3 DaT is present on one side only.
Remarks. Latzel (1880) and Verhoeff (1937) gave brief accounts of L.
tenebrosus but both these authors seem merely to have copied Meinert’s
description. Meinert described the female gonopods of this species as having
three pairs of spurs, so that its being based on a female of L. nigrifrons with a
supplementary medial spur on one gonopod only has never been suspected.
Latzel and Haase, in their original description of L. nigrifrons from various
parts of western Austria, stated that the structure of the 14th and 15th legs in
the male is variable, sometimes with flattening of the 15th femur and dorsal
sulci on the 14th and 15th tibiae, sometimes with only one or other of these
features and sometimes with neither. Loksa (1948) believed that these
TYPE SPECIMENS OF LITHOBIUS
25
characters are correlated with geographical distribution and he defined a new
subspecies, L. nigrifrons sulcatipes, to receive those individuals presenting
them. He (Loksa, 1955) separated the females of sulcatipes from those he
believed to belong to the typical form by means of their slightly longer genital
spurs and a difference in the number and arrangement of the setae on the
gonopod. The spurs on the gonopods of the holotype of L. tenebrosus (Fig. 11)
are intermediate in length between those figured by Loksa for L. nigrifrons
nigrifrons (Loksa, 1955: Fig. 38) and those he figured for L. nigrifrons
sulcatipes (Loksa, 1948: Fig. 1; 1955: Figs 39 & 40), and the arrangement of
the setae does not agree exactly with Loksa’s figures of either of these forms
(Loksa, 1955: Figs 38 & 39). I t seems likely that the subspecies sulcatipes
Loksa is without validity and that the sculpturing of the 14th and 15th legs in
the male varies quite independently of geographical distribution as Latzel and
Haase implied. The state of these characters in the typical L. tenebrosus (which
now becomes the correct name for this species) can only be determined by
examining males from the neighbourhood of Rasa, the type-locality.
Lithobius falteronensis Manfredi was based on a single male without
modification of the posterior legs from Mte. Falterona (Italy) which differs
from L. nigrifrons as originally described in spinulation, notably in the
complete absence of the spine series VpF and the presence of 1 5 DaP.
However, these variations in spinulation, the latter of which is found in the
holotype of L. tenebrosus, are unlikely to be of any taxonomic significance.
15. Lithobius gracilipes Meinert
Lithobius gracilipes Meinert, 1872: 3 18.
Type localities. Gerona, Seville (Spain).
Type specimens. Syntypes: two males, one immature, of L. rubriceps
labelled “Lithobius gracilipes Mein. Gerona, Sevilla M.”
Description. The adult is 22 mm long with 47 antennal articles, 1 + 13 ocelli,
6 + 6 prosternal teeth, traces of posterior projections on T.7, and 5 , 4, 4, 4
circular coxal pores. The immature specimen is 18 mm long with 48 antennal
articles, 1 + 9 ocelli, 6 + 6 prosternal teeth, no projections on T.7, and 4, 3, 3,
3, circular coxal pores. They are both conspecific with the syntypes of L.
insignis (see p. 22).
Remarks. Meinert distinguished this species from L. insignis by the fewer
ocelli, fewer coxal pores, and the absence of any ventral spine on the first
prefemur. But all these characters are such as would be expected in smaller
individuals which are not fully mature, or in adults which have retained certain
features of immaturity, and there is no justification for regarding the two forms
as taxonomically distinct: L. gracilipes thus becomes another synonym of L.
rubriceps Newport. This synonymy was, in fact, suggested very tentatively by
Meinert himself in his original description of the species and both Demange
(1958b) and Matic (Matic et al., 1967) suggested that insignis and gracilipes
might be no more than forms of the same species.
Brolemann (1924a), on the other hand, described a form from Morocco,
more slender, with fewer antennal articles, and with the tergites more rugose
than in the typical L. insignis, which he named L. insignis adrus. He later
E. H . EASON
26
(Brolemann, 1932) equated this form with L. graci2ipes Meinert. But neither of
the syntypes of gracilipes present these features and Brolemann’s form, if it is
indeed conspecific with L. insignis, should be known as L. rubriceps adrus
Brolemann.
16. Lithobiiis piilcher Meinert
(Fig. 12)
12
I
05
rnm
Figure 12. Lithobius forfiratus pulcher, dental margin of prosternurn, ventral.
Lithobius prdcher Meinert, 1872: 319.
Lithobirts Bonensis Meinert, 1872: 320.
T y p e locality Bbne (Algeria).
Tj.pe specimen. Holotype: a fairly well-preserved immature male of L.
forfi’catrts (Linn., 1758) labelled “Lithobius pulcher Mein. Bona M.”
Description. This specimen is 12 mm long with all the appearance of an
immature male (praematurus) of L. forficatus, the best known and most
widespread European species of Lithobius which has been fully described by
many authors. The dental margin of the prosternum, however, is much more
strongly recurved (Fig. 12) than in European specimens.
Remarks. Meinert distinguished this species from L. forficatus by its smaller
size, fewer ocelli, circular coxal pores, deficient ventral spinulation on the first
leg, and the absence of a second ventral spine of the 15th tibia. Brolemann,
who seems to have depended solely on Meinert’s description, included L.
piilcher in both his keys to north African species (Brolemann, 1924b, 1932). It
is surprising that the identity of this form, which is clearly conspecific with L.
forficatus and which, according to Meinert’s description, can only be separated
from it by features of immaturity, has not hitherto been recognized. However,
although the shape of the prosternum in a single individual might well be
attributable to aberration, the fact that all six type specimens of L. bonensis,
the next species to be considered, not only come from the same locality as the
holotype of L. prdcher and clearly belong t o L. forficatus, but also present the
same striking conformation of the prosternum, leads to the suspicion that all
these specimens belong to a distinctive geographical race. I t seems advisable,
therefore, to retain the name pulcher for an Algerian subspecies of L. forficatus
characterized by marked recurving of the prosternal dental margin.
17. Lithobius bonensis Meinert
Litfiobius Bonensis Meinert, 1872: 320.
TYPE SPECIMENS OF LITHOBIUS
27
Type locality. B6ne (Algeria).
Type specimens. Syntypes: five males and a female, all immature, of L.
forficatus pulcher labelled “Lithobius Bonensis Mein. Bona M.”
Description. These specimens, from 11.5 to 16 mm long, are all immature
examples (praematurus and pseudomaturus) of L. forficatus with the dental
margin of the prosternum strongly recurved as in the holotype of L. pulcher
(Fig. 12).
Remarks. The characters given by Meinert for this species indicate a degree
of maturity intermediate between that of the holotype of L. pulcher and an
adult of L. forficatus. Fedrizzi (1878) described some Italian specimens under
L. bonensis and Fanzago (1882) recorded this species from Italy, but Latzel (in
Dalla Torre, 1887) suggested that Fedrizzi’s description referred to L.
forficatus and Manfredi (1957) made the same suggestion in regard to
Fanzago’s record. Brolemann (1924b, 1932) included L. bonensis in both his
keys to North African species but seems to have depended solely on Meinert’s
description. The syntypes of L. bonensis all belong to the form which should
now be known as L. forficatus pulcher although most of them differ from the
holotype of pulcher in being rather more mature.
18. Lithobius grossidens Meinert
Lithobius grossidens Meinert, 1872: 324.
Type locality. Camorta, Nicobar Islands.
Type specimens. Syntypes: two females, one very mutilated, of L. pilicornis
Newport, 1844 labelled “Lithobius grossidens Mein. Kamorta Galatea”.
Description. These specimens have all the characters of L. pilicornis which
has been fully described by Eason (1964). Meinert was mistaken in recording
them as males.
Remarks. Latzel (1895) suggested L. grossidens as a possible synonym of L.
longipes Porat, 1870 (= L. pilicornis Newport). The Nicobar Islands are on an
ancient trade route and this common Mediterranean species must have been
introduced.
19. Lithobius rapax Meinert
(Figs 1 3 to 15)
Lithobius rapax Meinert, 1872: 325.
Lithobius stejnegeri Bollman, 1893: 199.
Lithobius sulcipes: Bollman, 1893: 199; Chamberlin, 191 lc: 260.
Monotarsobius arcticus Attems, 1909: 19.
Lithobius (Archilithobius) Haasei Attems, 1909: 22.
Ezembius stejnegeri: Chamberlin, 1919: 20.
Lithobius (Archilithobius 1 yamashinai Verhoeff, 1938: 103, Fig. 5.
Paobius haasei: Verhoeff, 1942b: 43.
Ezembius yamashinai: Chamberlin & Wang, 1952: 183.
Australobius yamashinai: Attems, 1953: 151.
? Lithobius sulcipes Stuxberg, 1876: 20.
E. H. EASON
28
? Lithobius ltaasei Garbowski, 1897: 248.
? Lithobius multiocellatiis Takakuwa, 1939: 33.
13
14
‘
05mrn
0 5mm
Figures 1 3 to 1 5 . Lirhobius rapax. 1 3 . Ocelli. 14. Dental margin of prosternum, ventral. 1 5 .
L,eft 15th metatarsus of d, medial (posterior).
Type locality. Sartung, Sakhalin (U.S.S.R.).
Type specimens. Lectotypes: a male with the 14th and 15th legs of one side
missing but otherwise fairly well-preserved, labelled “Lithobius rapax Mein.
Sartung 1869, Andria” is here formally designated as the lectotype. Paralectotypes; two rather mutilated males accompanying the lectotype and conspecific
with it.
Description of lectotype. Size: 19 mm long and 2.6 mm broad at T.lO.
Colour; dull yellow. Shape; breadths of head and of T.1,3, 5, 8,10,12 and 14
to each other as 61 : 57: 54 : 58 : 61 : 64 : 60 : 54.Head: broader than long;
lateral marginal interruptions distinct; posterior border feebly emarginate.
Antennae; 7.6 mm long; of 20 elongate articles. Ocelli: 1 + 5, 5, 5, 3 (Fig. 13).
Prosternum: with 3 + 3 teeth and a broad rounded median cleft; lateral spine
slender but much stouter than setae; lateral to the spine the anterior border is
almost straight, sloping gradually backwards (Fig. 14). Tergites: smooth with
some faint wrinkling; posterior border of T.l slightly emarginate, those of T.3,
5, 8, 10, 12, 14 and intermediate tergite moderately emarginate; posterior
angles of T.5, 8, 10 and 12 rounded, those of T.14 blunt, those of short tergites
all rounded without trace of projections. Coxal pores: 7, 7, 7, 5; large and
oblong except for medial pore on 12th t o 14th coxae which is small and
circular. Tarso-metatarsal articulations of anterior legs: very indistinct on first
three pairs, more distinct on 4th to 13th. 14th leg: stout relative t o anterior
legs but not obviously swollen: glandular pores less concentrated than in most
species of Lithobius: accessory apical claw well-developed; sensory spur very
long. 15th leg: 6.2 mm long; not obviously swollen; glandular pores as on 14th;
faint lateral sulci on femur and tibia and a faint dorsal sulcus on femur, all
quite inconspicuous; metatarsus stout throughout its length, not attenuated
distally as in most species of Lithobius so that its distal extremity is much
TYPE SPECIMENS OF LITHOBIWS
29
broader than the base of the apical claw which is somewhat uncinate; with a
small accessory apical claw; without sensory spur (Fig. 15); this unusual
structure of the metatarsus and apical claw may possibly be due to deformity
following abnormal regeneration. Chaetotaxy of legs: setae fairly long and
dense, particularly on the more anterior legs; the seriate arrangement of the
setae on the ventral aspects of the tarsus and metatarsus is quite clear on the
first thirteen pairs, rather indefinite on the 14th and absent on the 15th.
Genitalia: gonopod of a single article; second genital sternite without setae.
Spin ula t ion :
Ventral
1
2 -8
9-11
12
13-14
15
Dorsal
C
t
P
F
T
C
t
P
F
T
-
-
am
am
am
am
am
a
-
(m)
m
m
m
amp
amp
amp
amp
amp
am
-
-
p
p
mp
amp
amp
amp
amp
amp
amp
amp
amp
amp
ap
ap
P
p
a
ap
aP
ap
P
-
-
a
a
a
-
-
P
p
-
Several spines are duplicated. 9 , 10 and 11 VmT are present on one side only.
Description of puralectotypes. About 20 mm long and agreeing in all
essential characters with the lectotype, except that one specimen has deformed
forcipules and a small supplementary lateral prosternal tooth on one side in
addition to the usual 3 + 3. The 15th legs are missing in both specimens so that
the structure of the metatarsus and apical claw cannot be confirmed.
Remarks. Although Meinert’s description of this species is fairly adequate,
no suggestion as to its identity has ever been made. There is, however, little
doubt that L. rapax belongs to a widespread Japanese and North Pacific species
known to most authors as L. yamashinai despite the fact that there are two
older available names for the same species in addition to rapax. L. yarnashinai
Verhoeff was based on an immature male from Hokkaido (Japan) and has also
been recorded from the adjacent island of Sakhalin (Chamberlin & Wang,
1952). L. haasei Attems, which Verhoeff (1942b) proposed as the senior
synonym of yarnashinai, was based on a female from Bering Island which
Attems (1909) believed .to be identical with L. sibiricus Gerstfeldt as described
by Haase (1880). Garbowski (1897) had, unknown to Attems, already
proposed haasei as a new name for L. sibiricus (sensu Haase) but Chamberlin
(1923) questioned the identity of this form with L. haasei Attems: L. haasei
Garbowski is not, therefore, an objective synonym of L. haasei Attems and
may possibly refer to a different species. On the other hand Chamberlin
(1923), after examining a number of specimens of Lithobius from Bering Island
and other islands in or adjacent to the Bering Sea, concluded that L. haasei
Attems is conspecific with L. stejnegeri Bollman and Mono tarsobius arcticus
Attems, both originally described from Bering Island, and also with the
specimens which Bollman (1893) and he himself (Chamberlin, 1911c) had
recorded as L. sulcipes Stuxberg from Bering Island and St. Paul’s Island
(Alaska) respectively. Chamberlin (1923) was, however, uncertain of the
identity of the true L. sulcipes Stuxberg from northwest Siberia.
Although L. haasei Attems was described (like rapax and yamashinai) as
30
E. H. EASON
hwing 3 + 3 prosternal teeth, and the other forms from Bering Island as having
only 2 + 2, Chamberlin (1923) explained this as being due to the larger size of
the specimens on which Attems based L. haasei. Chamberlin also pointed out
that the smaller specimens which Attems would have placed in Monotarsobius
arcticus frequently have very indistinct anterior tarso-metatarsal articulations,
easily mistaken for the fused articulations characteristic of Monotarsobius. Yet
another species, L. mzi2tiocellatus Takakuwa from Urup (Kuril Islands) is
probably identical with L. rapax and was regarded by Verhoeff (1942b) as a
sub-species of L. haasei Attems, but it cannot be proposed as a synonym with
confidence.
L. rapax belongs to a fairly well-defined group of species which includes a
large proportion of the Lithobiidae occurring in eastern and northern Asia. This
group is characterized chiefly by the number of antenna1 articles being
restricted to 20 (19 to 21) and is distinguished from the predominantly
European Monotarsobius by larger size and the presence of tarso-metatarsal
articulations on the anterior legs of mature individuals, and from the southeast
Asian Aztstralobius by the different arrangement of the ocelli and the less
numerous prosternal teeth: it has as much claim to distinction as either
Monotarsobiits or Australobius, both of which are accepted by most authors as
either genera or subgenera of Lithobius.
Verhoeff (1942b) referred species of this group to Paobius which was
erected by Chamberlin (1916) to receive a number of western North American
species presenting several features (notably the absence of cephalic marginal
interruptions) not found in Asiatic forms. Ezembius, on the other hand, was
erected by Chamberlin (1919) as a monotypic genus to receive L. stejnegeri
Bollman so that Ezembius is the correct name for the group of Asiatic species
which can be arranged around L. rapax; but it should be regarded only as a
subgenus of Lithobitts.
It was, no doubt, Verhoeff‘s mention of 3 + 3 prosternal teeth in the original
description of L. yamashinai which induced Attems ( 1 95 3) to place this species
in Australobius. Species of Australobius do sometimes have only 3 + 3 teeth
but usually more and sometimes many more, whereas the species referable to
Ezembius usually have only 2 + 2 teeth and are distinct from those of
Airsfralobius in a number of other respects.
20. Lithobius suevicus Meinert
Lithobius suevicus Meinert, 1872: 326.
Type locality. Near Tubingen, Wurtemberg (Germany).
Type specimens. Syntypes: two males and a female of L. mutabilis L. Koch,
1862 labelled “Lithobius Sueviciis Mein. Tubingen M.”
Description. These specimens are identical with the lectotype of L. mutabilis
preserved in the British Museum (N.H.) (Reg. no. 13.6.8.435). They would
therefore run to L. mutabilis var. capathictis Verhoeff in Verhoeff‘s (193 5 ) key
to the infraspecific forms of this species (see Eason, 1972b). L. mutabilis has
been fully described by Matic. ( 1 966).
Remarks. Latzel (1880) recognized L. suevicus as a synonym of L. mutabilis
L. Koch and this synonymy has never been disputed.
TYPE SPECIMENS O F LITHOBIUS
31
2 1. Lithobius hispanicus Meinert
(Figs 16 to 21)
I
16
00
0
0
O0B0O
17
0.2
rnrn
0°80&
19
0.2 mrn
Figures 16 to 21. Lithobius hispanicus. 16. Ocelli of lectotype. 17 to 19. Ocelli of
paralectotypes. 20. Dental margin of prosternum, ventral. 21. Left 0 gonopod, ventral.
Lithobius Hispanicus Meinert, 1872: 327.
Lithobius inops Brolemann, 1932: 61.
Lithobius mundanus Verhoeff, 1937: 209, Figs 31 and 32.
Lithobius alicatai Matic in Matic et al., 1967: 182, Figs 1-4.
? Lithobius (Alokobius) canalensis Attems, 1952: 343.
Non Lithobius hispanicus: Attems, 1952: 345.
Type localities. Escorial; Granada; Seville; Caratraca near Malaga (Spain).
Type specimens. Lectotype: a well-preserved female selected by D. Schmidt
labelled “Lithobius Hispanicus Mein. Hispania M., Collin, L. Lund” is here
formally designated as lectotype. Paralectotypes: six females and three males,
some immature and all more or less mutilated, accompanying the lectotype and
conspecific with it, together with an immature male of L. castaneus Newport.
Description of lectotype. Size: 14.4 mm long and 1.6 mm broad at T.lO.
Colour: brown with an olivaceous tinge. Shape: breadths of head and of T . l , 3,
5, 8, 10, 12 and 14 to each other as 59 : 52 : 61 : 70 : 78 : 81 : 77 : 62. Head:
as broad as long; lateral marginal interruptions distinct; posterior border
straight. Antennae: 4.0 + 4.4 mm long; of 24 + 25 articles, about as broad as
long or slightly elongate. Ocelli: 1 + 3, 3, 1 (Fig. 16); organ of Tomosviry
small, adjacent to foremost ocelli, not remote. Prosternum: with 2 + 2 small
teeth and a broad shallow median cleft; lateral spine setiform (Fig. 20).
Tergites: T. 1 almost semi-circular with posterior border slightly convex;
posterior border of T.3 very slightly convex, that of T.5 very slight emarginate,
those of T.8, 10, 12, 14 and intermediate tergite slightly emarginate; posterior
angles of T.5, 8, 10 and 12 rounded, those of T.14 blunt, those of short tergites
E. H. EASON
32
all rounded without trace of projections. Coxal pores: 2, 2 , 2 , 2; circular.
Tarso-metatarsal articulatioris oj' anterior legs: rather indistinct on some legs.
14th leg: moderately swollen; accessory apical claw well-developed. 15th leg:
3.6 mm long; moderately swollen: accessory apical claw very small. Gonopod:
with 2 + 2 rather slender spurs; claw long and narrow with a distinct proximal
lateral denticle and a smaller distal medial denticle (Fig. 2 1).
Spin ula ti0 n :
Ventral
C
t
P
Dorsal
F
T
C
t
P
F
T
1
2
3
4-1 1
12
13
14
15
1 3 VaT
IS
present on one side only.
Description of adult paralectotypes. Differing from the lectotype in the
following characters: Antennae: of 18 to 27 articles, frequently different on
either side of the same individual. Ocelli: 1 + 4,3 , 2; 1 + 3 , 3, 2 or 1 + 3, 2, 1;
the arrangement can also be seen as two relatively large ocelli preceded by a
rosette, sometimes incomplete or distorted, of smaller ones (Figs 17 to 19).
Prosternum: with 2 + 3 teeth in one specimen. Female gonopod: spurs
sometimes larger than in the lectotype but of the same general shape; claw
either with a distinct proximal lateral denticle and a very feeble distal medial
denticle, or almost simple with a very feeble proximal lateral denticle only as
figured by Matic for L. alicatai (Matic, 1968: Fig. 2). Spinulation: the best
preserved specimen, a female 15 mm long with ocelli exactly as in the lectotype
and the genital claw almost simple, has the following:
Ventral
C
1
2
3
4-8
9
10-1 1
12
13
14
15
t
P
Dorsal
F
T
C
t
P
F
T
rn
m
(ah
am
am
am
am
am
m
m
3 VaF, 9 DaP and 2 DpF are present on one side only. 15 VaP is present o n one side of
another female.
The only male among the paralectotypes conspecific with the lectotype and
with the 14th and 15th legs intact is 12.5 mm long with 23 antenna1 articles,
1 + 3, 3 , 1 ocelli, 1, 2, 2, 1 coxal pores, spinulation very much as in the
lectotype, and six setae on either side of the first genital sternite. The 14th and
TYPE SPECIMENS O F LITHOBIUS
33
15th legs are more swollen than in females, particularly the 15th femur which
bears a very small nodular swelling at the distal extremity of its dorsal aspect;
the 15th tibia is flattened with a very shallow dorsal sulcus; the 15th accessory
apical claw is vestigial. This specimen may not be fully mature.
Remarks. Although there are a number of references to, and brief
descriptions of, this species in the literature it is not well-known and there has
been some confusion as to its identity. Attems (1903) described a single male
from Tunis very briefly: he probably had an example of L. hispanicus before
him when he wrote this description but he made no mention of the accessory
apical claw on the 15th leg which is very small and easily overlooked. Attems
(1927) repeated this omission in his key to the species of Archilithobius in
which hispanicus is described as having a simple 15th apical claw. Later the
same author (Attems, 1952) described an altogether different species from
Spain under L. hispanicus. Both Brolemann (1932) and Demange (1959)
discussed the uncertainty surrounding the identity of this species which has
arisen from Attems’s conflicting descriptions and Demange suggested that the
last one probably refers to L. pyrenaicus Meinert (see p. 41). None of the
characters given for L. hispanicus in the keys to Iberian and North African
species which have been published (Brolemann, 1924b, 1932; Demange, 1958b;
Matic et al., 1967) add anything to Meinert’s original description and the
species was first adequately redescribed by Machado (1952) from Portugal.
Matic (1968) described some Spanish specimens which he did not identify with
complete assurance but which almost certainly belong to L. hispanicus.
On the other hand two other species, L. inops Brolemann and L. alicatai
Matic, which cannot be separated satisfactorily from L. hispanicus, have been
described from Spain. Had it not been for Meinert’s mention of a third ventral
spine on the 15th prefemur of hispanicus (VaP which is present on one side
only of one paralectotype) Brolemann might never have named L. inops as a
separate species; and the characters used by Matic (Matic et al., 1967) to
separate L. alicatai from L. inops are unconvincing.
The spinulation of the legs, the arrangement of the ocelli and the shape of
the female genital claw have all been used as differential characters to separate
L. hispanicus, L. inops and L. alicatai from one another, but they are of no
value for this purpose. If the accounts of spinulation given for L. hispanicus by
Machado (1952), for L. inops by Brolemann (1949), Demange (1961a) and
Matic (1968), for L. alicatai by Matic (Matic et al., 1967; Matic, 1968), and for
the specimens doubtfully recorded by Matic (1968) as L. hispanicus are
compared with one another and with the spinulation of the lectotype and
paralectotypes of L. hispanicus it will be seen that they all conform to the same
fairly distinctive general pattern and the details do not suggest that more than
one species is involved. The arrangement of the ocelli in the type specimens of
L. hispanicus (Figs 16 to 19) can be seen as two separate ocelli preceded by a
rosette, as envisaged by Matic in L. alicatai (Matic et al., 1967: Fig. 2), or can
just as well be represented by a simple notation such as 1 + 3, 3, 2 which was
used by Machado (1952) to describe the ocelli in L. hispanicus. The female
genital claw is simple in L. inops as figured by Matic (1968: Fig. l), it bears a
very small lateral denticle in L. alicatai (Matic, 1968: Fig. 2), and in L.
hispanicus as figured by Machado (1952: Fig. XXIV, 2) and in Matic’s (1968)
doubtful examples of L. hispanicus this claw is tridentate as originally
3
34
E. H. EASON
described by Meinert. In the female type specimens of L. hispanicus the genital
claw varies in shape from almost complete simplicity with only a very small
lateral denticle, through claws with more prominent lateral and vestigial medial
denticles, to the frankly tridentate claw of the lectotype (Fig. 21).
The modification of the male 15th legs provides other characters used for
differentiating these three forms. Machado’s (1952), Brolemann’s (1932, 1949)
and Matic’s (Matic et al., 1967) descriptions of the femoral outgrowth in
hispanicus, inops and alicatai respectively, though different in detail, might all
refer to the same structure; and Demange’s (1961a) description of this
outgrowth in inops as “h peine marqtite”, its absence from Matic’s (1968)
doubtful examples of hispanicus, and its appearance in the male paralectotype
of L. hispanicus as an inconspicuous nodule might all be due to the immaturity
of the specimens. The shallow dorsal sulcus on the tibia of the male
paralectotype, also noted by Brolemann (1949) in inops, contrasts with the
small dorsal depression Matic found on the femur of alicatai (Matic et al.,
1967: Fig. 4); but these very feeble modifications are probably variable and are
most unlikely to be of taxonomic significance. Finally the remote position of
the organ of Tomosvary in alicatai (Matic et al., 1967: Fig. 2) and the third pair
of spurs on the gonopods of Matic’s single female of inops (Matic, 1968: Fig. 1)
could easily be due to individual variation and it seems certain that L.
hispanicus, L. inops and L. alicatai, all with a number of distinctive features in
common and all found in Spain, belong to single rather variable species.
The female from Portugal described by Verhoeff (1937) under L. mundanus
certainly belongs to L. hispanicus, and Attem’s (1952) description of L.
canalensis from Spain seems also to refer to this species but owing to its much
greater length (23 mm) it cannot be proposed as a synonym with confidence.
22. Lithobius lapidicola Meinert
(Fig. 22)
22
Figure 22. Lifhobius lapidicola, ocelli.
Lithobius lapidicola Meinert, 1872: 328.
Lithobius pusillus pitsillifrater Verhoeff, 1925: 152.
Non Lithobius lapidicola: Latzel, 1880: 106; Brolemann, 1930: 291 etc.
Type localities. Tubingen, Wiirtemberg (Germany). Rasa (South Tirol).
Roman Campagna; Sorrento (Italy) Cordoba; Granada (Spain). Bbne (Algeria).
Type specimens. Lecto type: a fairly well-preserved female selected by D.
Schmidt labelled “Lithobius lapidicola Mein. Bona Hispan. M. Italia Bgs.
Collin” and belonging to the form generally known as Lithobius pusillus
pusillifrater is here formally designated as the Iectotype. Paralectotypes: eight
females and four males accompanying the lectotype and conspecific with it,
together with seven males and four females of Turkobius sp., three females and
a male of L. apruzianus Verhoeff, two males and a female of L. boettgeri
Verhoeff, a male and a female, both immature, of L. erythrocephalus C. L.
Koch, a male of L. calcaratus C. L. Koch, a female of L. atrifrons Silvestri, a
female of L. margheritensis Verhoeff, a mutilated female of L. biporus
TYPE SPECIMENS OF LITHOBIUS
35
Silvestri? and seventeen mutilated specimens which could not be identified.
Description of lectotype. Size: 8.4 mm long and 1.05 mm broad at T.lO.
Colour; dull brown. Shape: breadths of head and of T . l , 3, 5 , 8 , 10, 12 and 14
to each other as 44 : 39 : 41 : 46 : 52 : 5 1 : 49 : 41. Head: slightly broader
than long; lateral marginal interruptions distinct; posterior border barely
emarginate, almost straight. Antennae: 2.4 mm long; of 30 articles, mostly
transverse. Ocelli: pale; 1 + 4, 2, 1 (Fig. 22). Prosternum: with 2 + 2 teeth;
lateral spine setiform; lateral to the spine the anterior border forms a feeble
shoulder. Tergites: moderately wrinkled and punctate; posterior borders of T. 1,
3 and 5 very slightly emarginate, those of T.8, 10, 22, 14 and intermediate
tergite moderately emarginate; posterior angles of T.5, 8 and 10 rounded, those
of T.12 blunt, those of T.14 angulated, those of T.9 obtuse, those of T . l l
squared, those of T.13 with traces of projections. Coxal pores: 2, 3, 3 , 3, and 2,
3, 4 , 3 ; circular; separated from one another by their own diameter or more.
Tarso-metatarsal articulations of anterior legs: quite distinct but with little
difference in breadth of articles. 14th leg: moderately swollen. 15th leg:
2.6 mm long: moderately swollen; accessory spical claw well-developed.
Gonopod: with two stout conical spurs; claw with a large distal medial denticle
almost as large as the tip of the claw itself and a much smaller more proximal
lateral denticle.
Spinulation:
Dorsal
Ventral
1
2
3 -8
9
10-11
12
13
14
15
C
t
P
-
-
-
-
-
-
-
-
m
m
-
F
T
C
t
-
-
-
am
am
amp
amp
amp
amp
m
m
m
m
m
m
m
m
-
-
-
-
mp
mp
mp
amp
amp
-
-
-
-
-
-
-
-
-
-
-
P
P
F
a
P
P
P
mP
mP
m P
m P
mP
p
aP
aP
aP
aP
P
P
P
-
T
a
aP
aP
aP
aP
P
-
Description of conspecific paralecto types. Differing from the lectotype in
the following characters: Length: 6.8 to 9.2 mm. Antennae: of 28 to 3 1
articles. Ocelli: 1 + 3, 2, 1 to 1 + 3, 3 , 2. Coxal pores: usually 2, 3, 3, 2.
Spinulation: the following is based on examination of the best preserved
specimens :
Ventral
C
t
P
Dorsal
F
T
C
t
P
F
T
36
E. H. EASON
Remarks. Of the syntypes, only those belonging to L. pusillus pusillifrater
and the single female of L. margheritensis answer at all closely to Meinert’s
description of L. lapidicola. The correct course, therefore, is to select one of
the former as lectotype since Meinert must have based his description to a very
large extent on examples of pusillifrater. His inclusion among the type
specimens of examples of larger species such as L. aprzizianus, which are up to
11 mm long, led to his giving the length of L. lapidicola as 8.5 to 11 mm:
pusillifrater rarely attains 10 mm in length so the identity of this species has
been overlooked. Meinert’s inclusion of so many examples of Tzcrkobius sp. is
altogether puzzling: they belong to a genus known only from the eastern
Mediterranean region (Chamberlin, 1952) and bear little resemblance t o L.
lapidicola.
The most widely accepted interpretation of Meinert’s description of this
species is that of Latzel (1880) which, as has already been explained (p. S),
refers to L. borealis. The forms described under L. lapidicola by Verhoeff
(1935, 1937) and Jeekel (1964a, 1964b), and also by Brolemann in his first
remark supplementary to his main description (Brolemann, 1930: 292), seem
to belong to yet another species or to a number of closely related species whose
identity is not fully understood. The true identity of L. lapidicola was,
however, suspected by Brolemann who, though following Latzel in his main
description of the species (Brolemann, 1930: 291), added a note in parenthesis
after his description of L. pusillus pusillifrater to the effect that this might also
be a form of L. lapidicola (Brolemann, 1930: 295).
Whether L. lapidicola (i.e. pusillifrater) is, in fact, conspecific with L.
pusillus Latzel as Verhoeff believed is questionable (Jeekel, 1964b)’ and it
should, for the time being, be regarded as a separate species. But there is no
doubt that, of the numerous infraspecific forms of L. pusillus which have been
described, L. pusillus denticztlata Attems (1904) from Dalmatia and L. pusillus
latialis Manfredi (1953) from the neighbourhood of Rome are either identical
with pusillifrater or much closer to it than to L. pusillits as originally described
by Latzel (1880): these two forms, if indeed they are subspecies, should
therefore be regarded as a subspecies of L. lapidicola Meinert and not of L.
p usill us Latzel.
Unfortunately, owing to Meinert’s method of labelling (see p. 2), it cannot
be said whether the lectotype of L. lapidicola comes from Italy, Spain or
Algeria; but pusillifrater was originally described from the Italian riviera ahd
has frequently been recorded from other parts of Italy so that Rome or
Sorrento are the most likely localities.
23. Lithobius eximius Meinert
Lithobius eximius Meinert, 1872: 3 3 3.
T y p e localities. Rasa (South Tirol). Roman Campagna; Ischia (Italy).
Granada (Spain). Constantine; Bbne (Algeria ). Tunis (Tunisia).
T y p e specimens. Syntypes: 1 5 females and 12 males, some immature, of L.
castaneus Newport, 1844 labelled “Lithobius eximius Mein. N. Africa M. Tirol
Ischia Collin.” A female, an adult male and an immature male of L. castaneus
labelled “Lithobius eximius Mein. Tunis M.”
TYPE SPECIMENS O F LITHOBIUS
37
Description. The adults are 19 to 27 mm long wirh antennae one third to a
half body length and coxal pores varying in shape from circular through oval
and oblong to frankly transverse. The female genital claw is simple in some
specimens (including the female from Tunis); in others it has a small lateral
denticle and in yet another the claw of the right gonopod is tridentate with
lateral and medial denticles while that of the left gonopod has a lateral one
only. In other respects the specimens agree with Rrolemann’s (1930)
description of L. castaneus.
Remarks. Pocock (1890) recognized L. eximius as a synonym of L. castaneus
Newport and this synonymy has never been disputed. The shape of the female
genital claw, which was described as simple in eximius by Meinert and is simple
in the holotype of L. castaneus from Sicily (Eason, 1972a), has been used by
Latzel (1880) and subsequent authors to distinguish between this and L. audax,
the next species t o be considered. In fact, the shape of this claw is variable in L.
castaneus, sometimes being different on either side of the same individual, and
is unlikely to be of much taxonomic value (see p. 38).
24. Lithobius audax Meinert
Lithobius audax Meinert, 1872: 334.
Type localities. Rasa (South Tirol). Gennazano; Rocca di Cavi (Italy).
Type specimens. Syntypes: six males and three females, some immature, of
L. castaneus labelled “Lithobius audax Mein. Tirol M. Italia Bgs.”
Description. The adults are 17 to 20 mm long with antennae one third of
body length and circular, oval or oblong coxal pores. The shape of the median
prosternal cleft and the size and spacing of the prosternal teeth all present
considerable variation. The genital claw is quite simple in one of the females,
simple with a feeble lateral lobe in another, and with a small sharp lateral
denticle in the third. In other respects the specimens agree with Brolemann’s
(1930) description of L. castaneus.
Remarks. Meinert distinguished L. audax from L. eximius (i.e. castaneus) by
its smaller size, shorter antennae, more circular coxal pores, and the shape of
the female genital claw which he described as “obsolete bilobus”, probably
taking this last character from the second of the female syntypes described
above. Latzel (1880) described L.audax as distinct from L. eximius, with the
genital claw “mit nur einem kleinen Seitenzahnchen (an der Aussenseitel”.
Subsequent authors (Brolemann, 1930; Demange, 1958b; Matic, 1959), in
addition t o the shorter antennae and the dentate female genital claw, used the
absence of the spine VaP from the foremost legs t o characterize audax which
they regarded as a subspecies of L. castaneus.
Owing to Meinert’s method of labelling (see p. 21, the single female of
eximius from Tunis (see p. 36) is the only female among the type specimens
of either eximius or audax from a known locality. This specimen together with
the holotype of L. castaneus from Sicily, the female syntype of L. Zucasi
Newport (= castaneus) from Algeria (see Eason, 1972a) and a number of
females of L. castaneus from Tunis and Algeria preserved in the British Museum
(N.H.) (Reg. nos. 1889.11.14.3; 1890.12.16.5.15; 1891.1.18.9) all have simple
genital claws with or without a feeble lateral lobe, so that it would be fair to
38
E. H. EASON
assume that a simple claw is always found in the form of the species occurring
in Sicily and North Africa were it not for Brolemann’s (1931) record of L.
~ “ r ~ audus
e t ~from Algeria, which suggests that a dentate claw may be found
in African examples. Latzel (1880), who admitted to being relatively unfamiliar
with L exirnius which he knew chiefly from Meinert’s description, recorded L.
a d a x from Tirol and much of Austria, so that the form of the species with a
dentate female genital claw is, presumably, predominant in this region. But of
two females from Liguria (northern Italy) preserved in the British Museum
(N.H.) (Reg. nos. 1889.3.8.24,26 (part) ), both of which clearly belong to L.
castuneus, the first has simple claws on both gonopods and the second has the
claw bidentate on one gonopod and simple on the other, so that it is difficult
to define two distinct subspecies on either geographical or morphological
grounds. Further, there is no correlation between the shape of the female
genital claw, the length of the antennae or the anterior extent of the spine VaP
in the specimens of L. custanerts which have been examined.
Even if the subspecies uudux (sensu Latzel) is valid, which seems unlikely, the
use of Meinert’s name for this form must now depend on the assumption that
he based his description of the female of L. utidax on the single syntype with a
dentate genital claw, and the further assumption that this specimen comes from
Rasa in South Tirol and not from either of the type localities in the Italian
peninsula where the form of the species with a simple claw is common if not
predominant (Verhoeff, 1934, 1942a), and where a female presenting a dentate
claw could be regarded as no more than a variant. Neither of these assumptions
seem to be justified and L. audux should be regarded as a synonym of L.
custaneus Newport.
Although Verhoeff (193 1) proposed L. audux as a synonym of L. castaneus
he seems to have based this synonymy on acceptance of Latzel’s (1880) use of
the name together with the incorrect assumption that a dentate female genital
claw is typical of L. custaneus. He referred examples of the species with a
simple genital claw, which undoubtedly belong to the typical form, either to L.
custuneus sangranus Verhoeff which he based on individuals from Molise (Italy)
with circular coxal pores and closely set prosternal teeth (Verhoeff, 1934), or
to L. custuneus buchnerorum Verhoeff, based on individuals from Ischia and
Sorrento (Italy) with oblong coxal pores and widely spaced prosternal teeth
(Verhoeff, 1942a). In fact, the shape of the coxal pores and the spacing of the
Drosternal teeth are no less variable in L. custuneus than the other characters
khich have been considered, and should not be used to define subspecies (see
also Matic, 1961).
25. Lithobius jlavtis Meinert
(Fig. 23)
23
0 5 mm
I
1
Figure 23. Lirhobius f l a w s (= inemis), dental margin of prosternum, ventral.
TYPE SPECIMENS OF LITHOBIUS
39
Lithobius j7avus Meinert, 1872: 336.
Type locality. Caratraca near Malaga (Spain).
Type specimen. Holotype: a badly mutilated male of L. inermis L. Koch,
1856 labelled “Lithobius j7uvus Mein. Caratraca L. Lund.”
Description. The specimen is so mutilated that the following description is
necessarily rather incomplete. Size: 15 mm long and 1.8 mm broad at T.8.
Colour: dull yellow. Shape: breadths of head and of T.l, 3, 5 , 8 , 10 and 1 2 t o
each other as 90 : 85 : 87 : 87 : 90 : 87 : 82. Head: broader than long; lateral
marginal interruptions distinct; posterior border barely emarginate, almost
straight. Antennae: about half body length; of 43 articles, equal in length and
breadth or somewhat transverse. Ocelli: obscured owing to moulting. Prosternum: with 3 + 3 teeth, the medial pair the larger and the line of their apices
very slightly procurved; lateral spine stout, dentiform, moderately pigmented;
immediately adjacent to the insertion of the spine the anterior border forms a
marked angle (Fig. 23). Tergites: T.l, 3 and 5 each with a pair of faint curved
paramedian sulci arising from the posterior marginal sulcus much as in L.
castaneus; posterior border of T. 1 barely emarginate, almost straight, those of
T.3 and 5 very slightly emarginate, those of T.8, 10 and 12 moderately
emarginate; T.14 and intermediate tergite crushed; posterior angles of large and
short tergites all rounded, without projections. Genitalia: crushed but
apparently immature.
Spinulation:
Dorsal
Ventral
1
2
3-8
9-12
13
14
C
t
P
T
C
-
-
mP
mP
mP
amp
amp
amp
am
am
amp
amp
am
a
a
-
m
m
t
P
F
T
-
-
-
-
amp
amp
amp
amp
amp
amp
ap
ap
aP
ap
p
P
a
ap
aP
ap
ap
P
-
Both the 14th and 15th legs are missing from this specimen but a single detached leg
without an accessory apical claw, from which the above spinulation of the 14th leg is
taken, seems to belong to the 14th pair. If the specimen had been similarly damaged when
examined by Meinert, he might have mistaken this leg for one of the 15th pair. Such a
mistake would explain his recording the ventral spinulation of the 15th leg, which is usually
1, 3 , 2 , 1 in L. inermis, as 1 , 3, 3 , 2 .
Remarks. Those authors who have included L. flavus in their keys to Iberian
species (Brolemann, 1924b; Demange, 1958b; Matic et al., 1967) have added
nothing t o Meinert’s description and no suggestion as to its identity has ever
been made. Failure to identify L. flavus with L. inermis L. Koch seems to have
been due to Meinert’s description of the former having been based on an
immature specimen with the detached 14th leg mistaken for the 15th.
L. Koch’s original description of L. inermis from near Malaga, the type
locality of L. flavus, is very scanty (Koch in Rosenhauer, 1856) but his
redescription (L. Koch, 1862) is recognizable. Brolemann (1930) redescribed
the species: he gave details of the spinulation of the legs but, like Koch, he
counted the stout lateral prosternal spine as a tooth making a total of 4 + 4,
and made no note of the details of sculpturing of the first three large tergites.
E. H. EASON
40
The lateral spine is, indeed, easily mistaken for a tooth (Fig. 23) and the
paramedian sulci on the tergites can only be seen on very careful examination.
Both Koch (1862) and Brolemann described the 15th leg with a ventral groove
and a medial concavity on the prefemur, a feeble dorsal sulcus on the femur,
and a deep medial sulcus on the tarsus and metatarsus, presumably in both
sexes. Attems (1952) described a new species, L psacadonotus. based on
several specimens from three separate localities in southern Spain, which almost
certainly belongs to L. inermis: he made no mention of any special sculpturing
of the 15th leg, but ventral and medial (posterior) features of the legs are not
usually distinctive in species of Littiobius and, being unexpected, are easily
overlooked on routine examination unless they take the form of projections:
the only dorsal feature noted by Koch and Brolemann, the feeble dorsal
femoral sulcus, is probably inconspicuous. Attems’s failure to mention these
modifications in his description of L. psacadonotus cannot, therefore, be
regarded as significant.
L castaneopsis Brolemann from Morocco (Brolemann, 1932, 1949) has most
of the essential characters of L. inermis including the paramedian sulci on the
anterior large tergites and the medial sulcus on the 15th tarsus and metatarsus.
The latter, although overlooked by Brolemann, was noted by Demange (1961a)
in both sexes. But Brolemann gave the number of antenna1 articles in
castaneopsis as 56 to 61 and described a dorsal sulcus on the male 15th tibia so
that this form cannot be identical with L. inermis and may represent a North
African subspecies. Demange (1961b) compared L. inermis, L. psacadonotus
and L. castaneopsis, which he regarded as separate species, and described a new
species, L. interruptus, based on a single apparently adult female from Minorca
which closely resembles L. inermis except for being only about half its length:
the exact status of L. inferrzcptus Demange can only be determined by
examining a male.
26. Lithobius gracilis Meinert
Lithobius gracilis Meinert, 1872: 3 37.
Non Lithobius gracilis Porat, 1869: 641.
Type locality. Escorial (Spain).
Type specimen. Holotype: a mutilated female of L. guadarramus Matic,
1968 labelled “Lithobius gracilis Mein. Escorial Collin.”
Description. This specimen, 13 mm long with short antennae of 34 + 39
articles, agrees with the original description of L. guadarramus (Matic, 1968:
118, Fig. 3a,b, and c). The characteristic broad ventromedial sulcus on the 15th
tibia, tarsus and metatarsus and the bidentate genital claw are quite distinct
despite the mutilation of the specimen.
Spinulation:
Dorsal
Ventral
C
t
P
F
T
C
t
P
F
T
mp
amp
amp
amp
amp
amp
am
am
am
am
am
a
-
-
mP
mP
m P
amp
aP
aP
P
p
aP
aP
1
-
9
-
-
13
1s
-
rn
-
The legs for which spines are not recorded are all missing.
a
P
-
TYPE SPECIMENS OF LITHOBIUS
41
Remarks. Brolemann (1924b) included L. gracilis in his key to the species of
Iberia and Morocco but he only made use of the characters mentioned by
Meinert, who overlooked the sulcus on the three distal articles of the 15th leg.
However, the holotype clearly belongs to L. gztadarramz~swhich was originally
based on several specimens from a number of localities in Spain (Matic, 1968).
The name gracilis is preoccupied by L. gracilis Porat, 1869, so that the correct
name for L. gracilis Meinert is L. gziadarrarnus Matic.
27. Lithobius pyrenaicus Meinert (Fig. 24)
”
24
Figure 24. Lithobius pyrenaicus, Ocelli.
Lithobius Pyrenaicus Meinert, 1872: 3 37.
Lithobius latebricola Meinert, 1872: 3 39.
Lithobius inermis pyrenaicus: Brolemann, 1926: 264.
? Lithobius hispanicus: Attems, 1952: 345.
Type locality. Near Gerona (Spain).
Type specimens. Lectotype: a well-preserved male selected by D. Schmidt
labelled “Lithobius Pyrenaicus Mein. Gerona M.” is here formally designated as
the lectotype. Purulectotype: a rather mutilated female accompanying the
lectotype and conspecific with it.
Description of lectotype. In general agreement with Brolemann’s (1926,
1930) descriptions of L. inermis pyrenaicus. Length: 20 mm. Antennae: of 47
strongly transverse articles. Ocelli: 1 + 4, 3, 3, 1 (Fig. 24). Prosternum: with
2 + 2 teeth, the line of their apices straight; lateral spine slender; adjacent to
the insertion of the spine the anterior border forms a rounded shoulder as in
the figured specimen of L. latebricola (Fig. 26). Tergites: posterior angles of
T.5, 8 and 10 rounded, those of T.12 and 14 blunt, those of T.9 and 11
rounded without projections, those of T.13 squared. Coxal pores: 4, 5 , 5 , 4;
circular. 14th leg: moderately swollen; tibia broadly sulcate dorsally; metatarsus feebly sulcate ventromedially. 15th leg: moderately swollen, the swelling
extending to tarsus and metatarsus; tibia broadly and deeply sulcate dorsally;
tarsus and metatarsus sulcate ventromedially as in the figured specimen of L.
latebricola (Fig. 27); without accessory apical claw.
Spinulation:
Dorsal
Ventral
C
t
P
F
T
C
t
am
am
am
am
am
am
am
a
a
-
a
a
-
a
-
~
1
2 -6
7-9
10
11-12
13
14
15
15 VaF is dulpicated on one side.
P
F
T
42
E. H. EASON
Description of paralectotype (female). With 36 + 41 antenna1 articles and
differing from the male in that the 14th and 15th legs are not generally swollen
nor are their tibiae sulcate; but the 15th tarsus and metatarsus are stout and
sulcate as in the male. The gonopod bears two fairly long conical slightly
unequal spurs and a short simple claw with a rounded tip.
Remarks. Although the type locality of L. pj’renuicus is Gerona in the
Pyrenees, Brolemann’s (1 926) opinion that pyrerzaicus represents a northern
subspecies of L. inernzis L. Koch cannot be upheld in view of the occurrence of
L. latebricola (=pyrenaicus) near Granada in southern Spain (see p. 44).
Brolemann’s opinion was based, to a large extent, on the striking modification
of the 15th tarsus and metatarsus which inermis and pyrenuicus share. But, in
addition to L. guadarramus (see p. 40) and the two doubtful species, L.
castaneopsis and L. interruptus (see p. 40), another species, L. schubarti
Demange from Spain, has been described with an almost identical modification
(Demange, 1959). It thus seems that a sulcate 15th tarsus and metatarsus,
which is found in both sexes, is characteristic not of a single species but of a
western Mediterranean group possibly allied to the North American genus
Kiberbius (Chamberlin, 1916), and there is every reason for believing L. inermis
and L. pyrenaicus to be specifically distinct.
28. Lithobius latro Meinert
(Fig. 25)
25
Figure 2 5 . Lithobius latro, ocelli.
Lithobius latro Meinert, 1872: 338.
Lithobius mutabilis latro: Verhoeff, 1900: 159.
Type locality. Rasa (South Tirol).
Type specimens. Lectotype: a well-preserved male labelled “Lithobius latro
Razzes M.” is here formally designated as the lectotype. Paralectotype; an
immature female accompanying the lectotype but not conspecific with it and
doubtfully identified as L. lucifiigw L. Koch.
Description of lectotype. In general agreement with Matic’s (1966) description of L. latro. Length: 9 mm. Antennae: almost half body length; of 30 + 3 3
articles. Ocelli: 1 + 4, 3, 2, 1 (Fig. 25). Prosternum; with 2 + 2 teeth; as figured
by Matic (1966: Fig. 65B). Tergites: posterior angles of T.5, 8. 10 and 12
rounded, those of T.14 blunt, those of T.9 and 11 without projections, those
of T.13 with very feeble projections; setae along posterior border of telson
unremarkable, not arranged in tufts. 14th leg: markedly swollen; tibia sulcate
dorsally with an illdefined setose boss medial to the distal end of the sulcus as
figured by Matic (1966: Fig. 65D). 15th leg: markedly swollen; tibia broadly
and deeply sulcate dorsally with a ring of setae, more numerous medially,
surrounding the sulcus; without accessory apical claw. Genitalia: gonopod
bearing three setae; second genital sternite without setae.
TYPE SPECIMENS OF LITHOBIUS
43
Spin ula tion :
Ventral
C
t
P
Dorsal
F
T
C
t
P
F
T
m
m
am
am
am
am
am
a
1
2
3-9
10-11
12
13
14
15
On the 15th leg DmP and DpP are of much the same length as VaP and VpP. On the 14th
leg VmT is relatively long.
Remarks. Latzel (1880) recognized L. latro as a distinct species but Verhoeff
(1900) and Brolemann (1930) regarded it as a subspecies of L. mutabilis L.
Koch. The latter view cannot be upheld since Matic (1966) has found the two
forms to be sympatric in Rumania.
The lectotype of L. latro does not agree with Verhoeff's (1937) definition of
the nominate subspecies or with any of the other subspecies of latro defined
either by him or later by Loksa (1962), which are separated from one another
by the relative length of certain spines on the 15th prefemur and 14th tibia, the
degree of modification of the male 14th tibia, the number and arrangement of
the setae on the posterior border of the telson, the number of setae on the male
gonopod and second genital sternite, and the number of antenna1 articles. If
these subspecies are considered valid a new name must be found for L. latro
latro (sensu Verhoeff), but they are probably based on variable characters not
correlated with geographical distribution and are better disregarded.
29. Lithobius latebricola Meinert
(Figs 26 and 27)
Lithobius latebricola Meinert, 1872: 339.
27
'
I 0 rnrn
c
i
I
I
Figures 26 and 27. Lithobius larebricola (= pyrenaicus). 26. Dental margin of prosternum,
ventral. 27. Left 15th tarsus and metatarsus, ventromedial (ventroposterior).
E. H. EASON
44
7 y p e loculitj’. Neighbourhood of Granada (Spain).
T j y e specimens. Syntj*pes: two males and a female, all fairly well-preserved,
and a male and female, both mutilated, of L . pj!reizaicw labelled “Lithobius
latebricola Mein. Granada M.”
Description. In general agreement with Brolemann’s (1926, 1930) descriptions of L. inerniis pyreizaicus and clearly identical with the type specimens of
L. pyrerzaicus (see p. 41). The prosternum, however, although of the same
general shape as in these specimens, shows some variation in the number and
size of the teeth. A male with 2 + 2 teeth has the lateral pair smaller than the
medial so that the line of their apices is procurved (Fig. 26): one femfile has
2 + 3 teeth and the other has the prosternum deformed with an enlarged
semicircular median cleft, an enlarged medial tooth on the left, and an enlarged
tridentate medial tooth on the right making an apparent total of 2 + 4 teeth.
The 14th and 15th legs of both sexes present the same modifications as in the
specimens of pyrenaicus and the 15th tarsus and metatarsus of a male is figured
(Fig. 27).
Spinulation: the following is based on the best preserved specimens.
Dorsal
Ventral
C
1
2
3 -6
7 -9
10-11
12
13-14
15
t
P
F
T
( a h p (a)m
amp am
amp am
amp am
amp am
amp am
amp am
a(mp) a
C
t
-
-
a
a
-
P
F
T
The letters in brackets indicate variable spines.
Remarks. Meinert recorded the dorsal sulci on the male 14th and 15th tibiae
of L. latebricola but he failed t o note the medial sulcus on the 15th tarsus and
metatarsus, so that the identity of this form with L. pyrenaicus has been
overlooked. Brolemann (1924b) included L. latebricola in his key to Iberian
and Moroccan species, but he only used some of the characters recorded by
Meinert and omitted any mention of the tibia1 sulci. Demange (1958b) and
Matic (Matic et al., 1967), probably following Brolemann, repeated this
omission. Verhoeff (1934) described a female from southeastern France under
L. latebricola but the dorsal spinulation he gave for the 15th leg and the
absence of any mention of modification of the 15th tarsus and metatarsus
suggest that this specimen may belong to some species other than L.
pyrenaicus: in Verhoeff‘s (1937) key L. latebricola seems to be based on the
same specimen. There are other references to L. latebricola Meinert in the
literature but none of them are accompanied by a description and it seems
unlikely that any of them refer to L. pyrenaicus.
30. Lithobius galatheae Meinert
Lithobius longipes: Meinert, 1872: 323.
Lithobius Galatheae Meinert, 1886b: 110.
TYPE SPECIMENS OF LITHOBIUS
45
Type locality. Madeira.
Type specimen. Holotype: a rather mutilated female of L. pilicornis labelled
“Lithobius Galatheae Mein. L. longipes Mein. olim. Madeira Galatea M.”
Description. This specimen has all the characters of L. pilicornis which has
been fully described by Eason (1964).
Remarks. Meinert (1886b) placed undue importance on such variable
characters as the degree of granularity of the tergites, the number of ocelli, and
the number of coxal pores as means of separating species. Consequently he
believed that the specimen from Madeira which he had tentatively but correctly
referred to L. longipes Porat in 1872 belonged to a new species which he
named L. galutheae. In fact, as Pocock (1891a) pointed out, Meinert was not
justified in making this distinction and longipes and galatheue are both junior
synonyms of L. pilicornis Newport.
3 1. Lithobius exiguus Meinert
(Figs 28 to 30)
28
1
0.2 mm
1
Figures 28 to 30. Paitobius exiguus. 28. Ocelli. 29. Dental margin of prosternum, ventrd. 30.
Left 0 gonopod, ventral.
Lithobius exiguus Meinert, 1886b: 110.
Paitobius exiguus: Chamberlin, 1922: 302.
? Lithobius cantabrigensis var. zinus Chamberlin, 191la: 41.
Non Lithobius exiguus: Chamberlin, 191la: 40; 1911b: 104.
Type locality. Near New York (U.S.A.).
Type specimens. Lectofype: a mutilated female selected by R. E. Crabill
labelled “Lithobius exiguus Mein. Skovene Omkr. New York L. Lund 27/69
Spec. orig. Mein” is here formally designated as the lectotype. Purulectotype: a
mutilated male accompanying the lectotype but not conspecific with it and
doubtfully referred to Tidubius Chamberlin.
46
E. H . EASON
Descriptioii of lectotype. Size: 7.4 mm long and 1.4 mm broad at T.lO; the
trunk is very contracted and when alive the length must have been about 9 mm.
Colour; dull yellow. Shape: breadths of head and of T . l , 3 , 5, 8 and 10 to each
other as 44 : 39 : 42 : 46 : 50 : 52; T.12 and 14 damaged and unsuitable for
measurement. Head: as broad as long; lateral marginal interruption distinct;
posterior border straight. Anrennae: 3.2 mm long; of 3 1 + 3 3 articles, about as
broad as long or somewhat transverse; terminal article elongate. Ucelli: 1 + 4, 4,
3 (Fig. 28). Prosfernitin: with 2 + 2 teeth, the line of their apices straight:
lateral spine setiform; lateral to the spine the anterior border is almost straight,
sloping sharply backwards (Fig. 29). Tergites: smooth with some faint
wrinkling; T. 1 with lateral borders strongly convex, converging posteriorly,
posterior border straight; posterior border of T. 3 barely emarginate, almost
straight, that of T.5 very slightly emarginate, those of T.8, 10, 12 and 14
slightly emarginate, that of intermediate tergite straight; posterior angles of
large tergites rounded, those of T.9 rounded without projections, those of T. 1 1
with feeble rounded projections, those of T. 1 3 with small triangular projections. Cosal pores: 2, 3, 3, 2; circular; separated from one another by their own
diameter. Tarso-metatarsal articidatiorzs of anterior legs: distinct. 14th leg:
missing. 15th leg: 2.6 mm long: accessory apical claw well-developed; sensory
spur vestigial. Gonopod: with two fairly long conical spurs; claw with medial
and lateral denticles distinct (Fig. 30); dorsolateral setae in a single row, three
on the second article and two on the third.
Spin ula ti0 11 :
Ventral
C
1
2 -9
11
12
13
15
t
P
Dorsal
F
T
C
t
P
F
T
-
m p
mP
amp
amp
amp
amp
a
aP
ap
a
aP
ap
aP
.-
-
m
m
-
-
p
mp
mp
mp
amp
a m
am
amp
amp
am
m
am
am
am
a
-
-
-
-~
-
a
m
-
-
a
-
P
P
p
P
-
Both the 10th and 14th legs are missing, but DaC is present on the 14th coxa. The
spinulation and other characters of the 15th legs are taken from detached legs which are
assumed to belong to the 15th pair owing to the absence of DpT, which is present o n the
14th but absent o n the 15th in the majority of species of Paifobiusdescribed b y Chamberlin
(1922).
Remarks. Chamberlin (1922) implied that his earlier records of L. exiguus
(Chamberlin, 1911a,b) from a number of localities in North America were
based on misdetermined specimens, and in referring the species to Paitobius
Chamberlin, which constitutes a distinctive genus of North American species
with a well-defined geographical distribution, he was solely dependant on
Meinert’s description: he was unable to identify it with any species known to
him, possibly because of Meinert’s understatement of the number of ocelli
(Meinert gave 7). The lectotype of L. exiguus, however, agrees in all essential
characters with Chamberlin’s (1922) detailed description of Paitobius (Tunabius) zinus which he designated as the type species of the subgenus Tunabius
Chamberlin, and had originally described as a variety (subspecies) of L.
cantabrigensis Meinert (Chamberlin, 191 la). But Dr. R. E. Crabill (in litt.)
TYPE SPECIMENS OF LITHOBIUS
47
believes that there may be a number of species of Paitobius in North America
yet to be described, and that they may be separable only by characters which
have not yet been fully recognized: the identity of exiguits with Paitobitis zinus
is therefore doubtful.
32. Lithobius h d i i Meinert
(Figs 31 to 3 3 )
/
32
0.2 m m
1
I
33
I
/
0.1m m
I
Figures 3 1 to 3 3 . Nmnpabius lundii. 3 1 . Ocelli. 32. Dental margin of prosternum, ventral. 3 3 .
Distal end of right 14th tibia of d, medial (posterior).
Lithobius Lundii Meinert, 1886b: 1 1 1.
Nampabius lundii: Chamberlin, 191 3 : 60.
Nampabius michiganensis Chamberlin, 1914: 302.
Non Lithobius lundii: Chamberlin, 191 la: 39.
Type locality. Near New York (U.S.A.).
Type specimen. Holotype: a male labelled “Lithobius Lundii Mein. Skovene
om New York L. Lund.” Dr R. E. Crabill has attached a label to this specimen
indicating that he believed it might be identical with Nampabius michiganensis
Chamberlin.
Description. Size: 6.8 mm long and 1.2 mm broad at T.8. Colour: brown.
Shape: breadths of head and of T . l , 3, 5 , 8, 10, 12 and 14 to each other as
43 : 37 : 42 : 47 : 5 1 : 50 :48 : 40. Head: slightly broader than long; lateral
marginal interruptions distinct; posterior border straight. Antennae: 2.3 mm
long; of 20 articles, somewhat elongate distally. Ocelli: 1 + 3, 3, 2 (Fig. 31).
Prosternum: with 2 + 2 teeth, the line of their apices moderately recurved;
lateral spine setiform (Fig. 32). Tergites: smooth with some sparse and feeble
puncta; posterior borders of T . l , 3, 5 , 8, 10 and 12 slightly emarginate, that of
T.14 convex, that of intermediate tergite straight; posterior angles of T.5 and 8
rounded, those of T.10 and 12 blunt, those of T.14 rounded, those of T.9
E. H. EASON
48
obtuse, those of T . l l squared, those of T . 1 3 with small projections. Coxal
pores: 2, 3, 3, 2 ; circular; separated from one another by up to twice their own
diameter. Tarso-metatarsal artidations of anterior legs: very indistinct on
foremost legs; distinct on 12th. 14flz leg: 2.25 mm long; moderately swollen;
the dorsomedial aspect of the distal end of the tibia bears a small wart-like
outgrowth (Fig. 3 3 ) . 15th leg: 2.45 mm long; moderately swollen; accessory
apical claw present. Genitalia: gonopod of a single article; second genital
sternite without setae.
Spin u la t ion :
Dorsal
Ventral
C
t
P
F
T
C
t
P
F
T
Both the 2nd and 13th legs are missing
Remarks. Chamberlin (1913), in his original account of Nampabius, referred
L. lundii t o this distinctive and well-defined genus on the basis of Meinert’s
description, but he was unable to identify it with any species then known to
him. He subsequently (Chamberlin, 1914) described N. michiganensis from
numerous specimens, all taken near Douglas Lake, Michigan. There is little
doubt that the holotype of L. Ittndii belongs to michiganensis which differs
from the other species of Nampabius in presenting an enlarged posterior
isolated ocellus (Fig. 3 1 ) as in most species of Lithobius. Earlier records of L.
lundii from the Southern States of U.S.A. (Chamberlin, 1911a) were almost
certainly based on misdetermined specimens.
ACKNOWLEDGEMENTS
I wish to thank the Zoological Museum, Copenhagen University, and Dr S. L.
Tuxen for allowing me t o borrow the type specimens of Meinert’s species, and
Mr K. H. Hyatt of the Arachnida and Myriapoda Section of the Department of
Zoology of The British Museum (N.H.) for his help in enabling me t o examine a
number of specimens held by that Museum. My special thanks are due to Mr
Henrik Enghoff of Copenhagen University without whose personal help this
study could not have been undertaken.
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