zoological Journal ofthe Linnean Society (1982). 74: 9-33 . With 12 figures
A review of the north-west European species
of Lithobiomorpha with a revised key to
their identification
E . H . EASON
Bourton Far Hill. Moreton in Marsh.
Gloucestershire. GL56 91’.
England
Accepted for publication March 1981
All the species and subspecies of Lithobiomorpha known to occur in Austria. Germany. Switzerland.
Scandinavia. the Netherlands. Belgium and northern and central France are reviewed and a key to
their identification is given . Lithobius pypaeus Latzel and L . lusitanus valesiacus Verhoeff are fully
described. and references to adequate descriptions of all the other species and subspecies are given
among the synonyms . Further synonyms are given. where necessary. in order to resolve the confusion
which exists over nomenclature. Archilithobius stramineus Attems. 1927 is proposed as a junior synonym
of L . lusitanus Verhoeff. 1925. and L .pusillus Latzel. 1880 as ajunior synonym o f L. lapiduola Meinert.
1872.
KEY WORDS:-Lithobiidae
-
Henicopidae
-
north-west Europe
-
distribution
-
key to species.
CONTENTS
Introduction . . . . . . . .
Systematic review of species
. . . .
Family Lithobiidae
. . . . .
Genus Eupolybothrur Verhoeff . . .
Eupolybothrus grossipes (C. L . Koch) .
Eupolybothrus tridentinus (Fanzago) .
Genus Harpolithobius Verhoeff
. .
Harpolithobius anodus (Latzel) . .
Genus Lithobius Leach . . . .
Lithobius validus Meinert . . .
Lithobius forfrcatus (LinnC) . . .
Lithobius piceus L . Koch
. . .
Lithobius pilicornis Newport . . .
Lithobius castaneus Newport . . .
Lithobius dentatus C. L . Koch . .
Lithobius nodulipes Latzel . . .
Lithobius tenebrosus Meinert . . .
Lithobius macilentus L . Koch
. .
Lithobius pelidnus Haase
. . .
Lithobius cyrtopus Latzel
. . .
Lithobiur mutabilis L . Koch . . .
+
0024-4082/82/010009 25$02.00/0
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0 1982 The Linnean Society of London
10
E. H. EASON
Lithobiuc latro Meinert . . , . ,
.
Lifhobiuspygmaeus Latzel . . . . .
Lithobius subtilis Latzel.
. . . . .
Lithobius muficus C. L. Koch . . . .
Lithobius ralcarafus C. L. Koch . . ,
,
Lifhobius lurifugus L. Koch.
.
.
.
.
Lithobius melanops Newport.
. , . ,
Lithobius bicuspis Meinert . . . . .
Lithobius agilis C . L. Koch . . . . .
Lifhobius ervthrocephalus C . L. Koch . . .
Lithobius etythrocephulus schuleri Verhoeff . .
Iifhobiuc lucitanus VerhoeK.
. . . .
Lithobius lusiknus ualesiarus Verhoetr
. .
Lithobiuc borealis Meinert . . . . .
Lithobiuc lapidicola Meinert.
.
.
.
.
Lilhobius mirrops Meinert . . . . .
Lithobius (Monofarsobiuc! curtipes C. L. Koch .
Lithobius (Monotarsobius) crassips L. Koch .
Lithobius (A4onotarsobiusi aeruginosuc L. Koch.
Lithobius (hfonofarsobiuc!aushiacus (Verhoefn
Family Henicopidae . . . . . . .
Genus Lamycks Meinert ,
. . , . .
Lamyctes fulcirornis Meinert.
. . . .
Genus Lamyctinus Silvestri
. . . . .
I.amyctinus coeculus (Brolemann. . . . .
Key to the north-west European Lithobiomorpha .
Acknowledgements.
. . . . . . . .
References.
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31
INTRODUCTION
There has been considerable demand in recent years for a means of identifying
the species of Lithobiomorpha to be found in north-west Europe and in the present
paper an attempt is made to meet this demand. Most workers are using Verhoeffs
( 19373 comprehensive key to the European and Mediterranean Lithobiidae which
includes a large number of species, subspecies and varieties based on minor
variations and abnormal individuals which are not worthy of taxonomic status. A
more useful key of Verhoeff s for workers in north-west Europe, unencumbered by
redundant taxa, is restricted to those species of Lithobiomorpha occurring in
Germany and Switzerland Werhoeff, 1936): but some of the characters used in
this key are unreliable and consequently some of the species included are difficult
to recognize. Of the standard monographs, Latzel’s (1880b) account of the
centipedes of the Austro-Hungarian Empire still contains some of the best
published descriptions of many European Lithobiidae but it has very few
illustrations and gives no details of the spinulation of the legs: Latzel recorded
these spines numerically on the first, fourteenth and fifteenth legs only.
Brolemann’s ( 1930) monograph of the French centipedes includes descriptions of
the majority of north-west European species, some based on his own observations,
others copied from Latzel. Most of those omitted or inadequately described by
Brolemann are fully described and illustrated by Matic (1966) in his monograph of
the Rumanian anamorph Chilopoda. All the above descriptions applying to northwest European species are cited in the present paper among the synonyms, which
are designed to give references to adequate published accounts of each species and
to resolve the confusion which exists over nomenclature. Past records in the
literature of such species as Lithobius borealis Meinert, L. lapidicola Meinert and L.
mirrops Meinert, whose true identity has only recently been established, are
NORTH-WEST EUROPEAN LITHOBIOMORPHA
11
valueless unless accompanied by some description. Where none of the standard
monographs gives an adequate account of a species, reference is also made among
the synonyms to other more detailed published descriptions. No adequate
descriptions of L. pygmaeus Latzel or L. lusitanus valesiacus Verhoeff have been found
in the literature and both these forms are fully described.
No attempt has been made to give details of the known distribution of each
species in north-west Europe which is defined here to include Austria, Germany,
Switzerland, Scandinavia, the Netherlands, Belgium and northern and central
France, excluding the Pyrenees and the Mediterranean region where the lithobiid
fauna is markedly different from that of north-west Europe. But the notes on
distribution given by Latzel (1880b), Rothenbuhler (1899, 1901), Brolemann
(1930), Verhoeff (1935, 1936) and Jeekel (1964, 1977) have been consulted
together with Andersson’s (1979a) work on the Swedish Lithobiomorpha and some
unpublished records in the author’s possession, and a general statement is made in
regard to the distribution of each species. All the species known to occur in both
France and the Netherlands are likely to be found in Belgium for which there are
very few authentic records.
SYSTEMATIC REVIEW OF SPECIES
Family Lithobiidae
Genus Eupolybothrus Verhoeff
Eupolybothrus grossipes (C. L. Koch)
Lithobius grossipes C. L. Koch, 1847: 146; Latzel, 1880b: 44, figs 26, 27.
Bothropolysfasciatus: Brolemann, 1930: 245, fig. 389 (non Newport, 1844).
Polybothrusfasciatus: Verhoeff, 1936: 106; 1937: 179, figs 2, 5, 6, 7 (oben).
Eupolybothrus fasciatus: Matic, 1966: 66, fig. A-D.
Eupolybothrus grossipes: Eason, 1970a : 295, figs 4-7.
In their tables of the spinulation of the legs of this species Brolemann gave VpT,
VaTa and VpTa, and Matic gave VpT, on many of the legs. But none of these
spines has been found in the specimens examined for the purpose of the present
study and it has been suggested (Eason, 1970a : 290) that the enlarged spinous
setae often found in these positions have sometimes been mistaken for spines.
Eupolybothrus grossipes is known from Austria, southern Germany and
Switzerland.
E u p o ~ b o t ~ r utriden~inus
s
(Fanzago)
Lithobius tridentinus Fanzago, 1874: 36; Latzel, 1880b: 49.
Lithobius terreus Fedrizzi, 1877: 203 ; Latzel, 1880b: 56.
Lithobius leptopus Latzel, 1880b: 53.
Polybothrus leptopus: Verhoeff, 1936: 106; 1937: 178, fig. 10.
Eupolybothrus leptopus: Matic, 1966: 7 1, fig. 27 A-D.
Eupolybothrus tridentinus: Eason & Minelli, 1976: 186, fig. 1.
Latzel, when describing L. tridentinus and L. terreus as distinct from L. leptopus,
was merely copying earlier descriptions of these species, both of which gave undue
weight to variable characters. Eason & Minelli (1976) have examined the holotype
of L. tridentinus and have shown that it is identical with both L. terreus and L.
leptopus.
I-'
E. H . EASON
Eupolybothrus tridentinus is known from i2ustria, southern Germany and
Switzerland.
Genus Harpolithobius Verhoeff
Harpolithobius anodus (Latzel)
Lithobius anodus Latzel, 1880b: 88.
Harpolithobius anodus: Verhoeff, 1936: 102; 1937: 237; Matic, 1966: 78, fig. 30
Ii-G.
Harpolithobius anodus is known from -4ustria.
Genus Lithobius Leach
Lithobius validus Meinert
Lithobius validus Meinert, 1872: 291 ; Latzel, 1880b: 50, figs 14, 25, 28; Verhoeff,
1936: 103; 1937: 196.
Lithobiuspunrtulatus: Brolemann, 1930: 250; Matic, 1966: 110, fig. 40 A, B (non C.
L. Koch, 1847).
Lithobius validus is known from Austria, southern Germany and Switzerland.
Lithobius fotjicatus (Linne)
Srolopendraforjrata Linne, 1758: 638.
Lithobius forJiratus: Latzel, 1880b: 57; Brolemann, 1930: 257, figs 378-380,
393-397; Verhoefl', 1936: 103; 1937: 198; Matic, 1966: 118, fig. 45 A-C.
Lithobius forjratus is known from Austria, France, Germany, the Netherlands,
Scandinavia and Switzerland.
Lithobius pireus L. Koch
Lithobius pireus L. Koch, 1862: 49, fig. 15; Latzel, 1880b: 64; Brolemann, 1930:
260, fig. 377; Verhoeff, 1936: 103; 1937: 199; Matic, 1966: 125, fig. 47 A-E.
Lithobius pireus gracilitarsis Brolemann, 1898: 200; 1930: 262, figs 398-401 ;
Verhoeff, 1937: 199.
Brolemann (1898) named a Pyrenean form with long slender 15th tarsi and
metatarsi as a distinct subspecies, grarilitarsis, but Eason (1972) has shown that the
typical form of the species frequently has this character. Females ofL. piceus usually
have a bidentate claw on the gonopod: those found in Switzerland with a simple
claw are regarded by Demange (1958) as belonging to a distinct subspecies, L.
pireus verho@ Demange. Although usually with 4 + 4 or more prosternal teeth,
examples of L. pireus may have only 4 3 or 3 3 teeth and those with the latter
number are keyed separately.
Lithobius pireus is known from Austria, France, Germany, the Netherlands and
Switzerland.
+
+
Lithobius pilirornis Newport
Lithobius pilirornis Newport, 1844: 96; Brolemann, 1930: 297, figs 373, 375, 376,
381, 384; Verhoeff, 1937: 203.
Lithobius pilirornis is known from a few Atlantic coastal areas in France and the
Netherlands.
NORTH-WEST EUROPEAN LITHOBIOMORPHA
13
Lithobius castaneus Newport
Lithobius castaneus Newport, 1844: 96 ; Brolemann, 1930: 3 1 1, figs 439-44 1 ;
Verhoef€, 1937: 205.
Lithobius eximius Meinert, 1872: 333; Latzel, 1880b: 123.
Lithobius audax Meinert, 1872: 334; Latzel, 1880b: 124.
Lithobius castaneus var. audax: Brolemann, 1930: 314.
Some authors regard audax, which has a dentate as opposed to a simple claw on
the female gonopod, as a variety or subspecies of L. castaneus or even as a distinct
species. There does seem to be a tendency for a dentate claw to occur more
frequently in northern examples of L. castaneus than in those from Sicily (the type
locality) and North Africa, but it is difficult to draw any clear distinction between
the two forms (Eason, 1974). Although massive porodonts and small posternal
teeth are usually found in this species, some specimens have relatively slender
porodonts, others have quite well-developed teeth (Eason, 1980a: fig. 1) : the
latter are keyed separately.
Lithobius castaneus is known from southern Austria.
Lithobius dentatus C. L. Koch
Lithobius dentatus C. L. Koch, 1844: 22, fig. 22; Latzel, 1880b: 81 ; Brolemann,
1930: 263; Verhoeff, 1936: 103; 1937: 202; Matic, 1966: 51, fig. 58 A-D.
Lithobius dentatus is known from Austria, Belgium, France, Germany, the
Netherlands and Switzerland.
Lithobius nodulipes Latzel
Lithobius nodulipes Latzel, 1880a, 225; 1880b: 68, figs 30, 30a; Verhoeff, 1936: 103;
1937: 200, figs 23, 36, 37; Matic, 1966: 134, fig. 51 A-D.
Lithobius nodulipes is known from Austria, southern and eastern Germany and
Switzerland.
Lithobius tenebrosus Meinert
Lithobius tenebrosus Meinert, 1872: 317; Latzel, 1880b: 70; Verhoeff, 1937: 201.
Lithobius nigrijirons Latzel & Haase in Latzel, 1880b: 71; Brolemann, 1930: 271;
Verhoeff, 1936: 103; 1937: 201; Matic, 1966: 146, fig. 56 A-C.
The original description of L. tenebrosus was based on a female with 3 2 spurs on
the gonopods instead of the usual 2 + 2 (Eason, 1974: fig. l l ) , which Meinert
described as having 3 + 3 such spurs. Consequently both Latzel and Verhoeff
( 1937) described tenebrosus and nigrqrons as separate species.
Lithobius tenebrosus is known from Austria, Germany, Scandinavia and
Switzerland.
+
Lithobius macilentus L. Koch
Lithobius macilentus L. Koch, 1862: 63, fig. 25.
Lithobius aulacopus Latzel, 1880b: 84; Brolemann, 1930: 266, figs 382, 404, 405;
Verhoef€, 1936: 104; 1937: 202; Matic, 1966: 142, fig. 55 A-E.
Lithobius rnacilentus is known from Austria, France, Germany, the Netherlands,
Scandinavia and Switzerland.
I4
E. H. EASON
Lithobius pelidnus Haase
1,zthobius pelidnus Haase, 1880: 33; Latzel, 1880b: 95; Brolemann, 1930: 306;
I’erhoeff, 1936: 104; 1937: 69; Matic, 1966: 170, fig. 67 A-D.
Lithobius pelidnus is known from every country in North-west Europe.
Lithobius cyrtopus Latzel
Lithobius cyrtopus Latzel, 1880a: 225; 1880b: 93; Verhoeff, 1936: 104; 1937: 205;
Matic, 1966: 167, fig. 66 A-D.
Most examples of this species have a 15th accessory apical claw but some
individuals, usually males, are without this claw and are keyed separately.
Although Latzel ( 1880b) did not record L. cyrtopus from Austria as defined by its
present boundaries it is known from southeastern Germany.
Lithobius mutabilis L. Koch
Lithobius mutabilis L. Koch, 1862: 75, fig. 34; Latzel, 1880b: 97; Broleman, 1930:
304; Verhoeff, 1936: 105; 1937: 207; Matic, 1966: 161, fig. 63 A-D.
Lithobius mutabilis is known from Austria, France, Germany and Switzerland.
Lithobius latro Meinert
Lithobius latro Meinert, 1872: 338; Latzel, 1880b: 102; Verhoeff, 1936: 105; 1937:
207; Matic, 1966: 165, fig. 65 A-D.
Lithobius mutabilis latro: Brolemann, 1930 : 306.
Lithobius latro is known from Austria, southern Germany and Switzerland.
Lithobius pygmaeus Latzel
(Figs 1-7)
Lithobius pygmaeus Latzel, 1880b: 86; Verhoeff, 1936: 104; 1937: 202.
Non Lithobius pygmaeus Sseliwanoff, 1880.
Latzel mentioned small posterior projections on T.7 in males of this species from
Austria, in addition to projections on T.9, 11 and 13 in both sexes. The Swiss
specimens described below differ from those of Latzel in having a larger average
number of antenna1 articles (Latzel gave 29-35) and in being without posterior
projections on T.7 and 9; but since they agree closely with Latzel’s description in
other respects, they undoubtedly belong to L. pygmaeus which is unique among
European species of Lithobius in combining small size with distinct tarsal
articulations on all the legs and well-marked dorsal sulci on the male 14th and 15th
tibiae. Although Rothenbuhler ( 1899) recorded L. pygmaeus, presumably the
typical form, from the Rhone valley in Valais, Switzerland, the present specimens
may belong to a distinct subspecies and are keyed separately. The species is also
known from eastern Germany.
Lithobius pygmaeus Sseliwanoff refers to a species not found in north-west Europe.
Material examined
Nineteen specimens from Switzerland collected by Dr Hans Jungen and his
students: three males and four females from the Schlosswald near Salez; two males
15
NORTH-WEST EUROPEAN LITHOBIOMORPHA
2
/
TO
TO
TO
4
L
I
0.1mm
I
I
0.1m m
0.1m m
Figures 1-7. Lithobiwpygnaeus. Figs 1-3, right ocelli and organ ofTomosv6ry (TO); Fig. 4, prosternum,
dorsal; Fig. 6, left 0 gonopcd, ventral; Fig. 7, left 9 gonopod,
ventral; Fig. 5, right 15th leg of
dorsal.
a,
and three females from the ruins of the castle Furstenstein on the Blauen, Ettingen;
two males and a female from Suret between Aarau and Rupperswil; two males
from Eischberg, south of Aesch; a male and a female from Ramsflue near
Erlinsbach.
Description
Size: 5.8-7.8mm long and 0.78-1.04mm broad at T.lO. Head: 0.70-0.86mm
broad; about as broad as long, broader than T.3 but usually narrower than T.5;
E. H . EASON
16
no median thickening of posterior marginal ridge ; posterior border straight ;
paired posterior depressions usually well-marked. Antennae : two-fifths to almost
half body length; of 34-42 (usually 35-38) articles, as broad as long or transverse,
the last slightly to moderately elongate. Ocelli: 7-9 on each side; posterior ocellus
usually rather smaller than posterosuperior ; 1 3, 2, 1 ; 1 3, 2, 2 and 1 3, 2, 2, 1
are the usual arrangements (Figs 1-3). Organ ofTomosvary : almost as large as the
largest ocellus (Figs 1-3). Prosternum: with 2 + 2 (rarely 2+3) teeth, the line of
their apices straight or very slightly recurved; porodont almost twice as broad as a
large seta at base, setiform at apex, arising close to the lateral tooth and usually
directed medially ; no shoulder lateral to porodont (Fig. 4). Tergites : finely rugose
with very little wrinkling; T. 1 narrower than T.3, approximately rectangular or
almost semicircular with posterior border straight; posterior borders of T.3 and 5
very slightly emarginate, those of T.8, 10 and 12 slightly emarginate, that of T.14
almost straight; posterior angles of T.7 obtuse, those of T.8 evenly rounded, those
of T.9 squared or with traces of projections, those of T . 10 abruptly rounded, those
of T.12 and 14 blunt, those of T . l l and 13 with short broad projections.
Intermediate tergite : posterior border moderately emarginate in females, strongly
so and frequently lobed at either angle in males. Coxal pores: 2, 3, 3, 2 or 2, 3 , 3, 3
in males; 2, 3, 3, 3-3, 4,4, 3 (frequently 2, 3, 4, 3) in females; circular, separated
from one another by their own diameter. Tarsal articulations of anterior legs :
distinct. 14th and 15th legs: tibia and femora moderately thickened in females,
usually more so in males; accessory apical claw two-thirds as long as principal claw
on both legs; 15th a third to two-fifths of body-length. Sensory spurs: longer than
accessory apical claw on first to 13th legs, half as long on 14th, very short on 15th.
Male secondary sexual characters: 14th and 15th tibiae each with a broad dorsal
sulcus extending over most of its length (Fig. 5) ; 13th tibia usually with a similar
but fainter sulcus. Glandular pores: concentrated on 14th and 15th legs only. Male
genitalia: second genital sternite with two setae; gonopod of a single broad article
with two setae. Female gonopod : with two short stout spurs and a claw with a welldeveloped medial denticle and a smaller lateral denticle (Fig. 6) ; one female has a
third spur on one gonopod; four or five stout dorsolateral setae in a line on the
second article, the largest almost twice as broad as a large general seta; three or
four very short and slender dorsomedial setae on the first article immediately
proximal to the insertions of the spurs (Fig. 7 ) .
+
Spinulation :
C
1
2
3
4
5
6
-
.-
-
7
8
9-10
I1
12
13
14
15
~
-
-
t
P
-
-
Ventral
F
+
+
Dorsal
T C
t
P
F
T
a
a
NORTH-WEST EUROPEAN LITHOBIOMORPHA
17
Letters in brackets indicate the usual variable spines. In addition to these
variations 11 VpP, 1 DaF, 5 DpF and 12 DaT are each present in one specimen
only; and 8 VaF, 12 VaF, 3 DaF and 12 DpF are each absent in one specimen
only.
Lithobius subtilis Latzel
Lithobius subtilis Latzel, 1880b: 91 ; Verhoeff, 1936: 104; 1937 : 206; Eason, 1980b:
125, figs 1-6.
Lithobius siluaenigrae Verhoeff, 1935: 186; figs 2, 3; 1937: 202.
? Lithobius lapidicola: Jeekel, 1964: 135 (non Meinert, 1872).
Most examples of this species are without distinct posterior projections on T.9
but some individuals have quite well-developed projections on this tergite and are
keyed separately.
Lithobius subtilis is known from Austria, Germany, the Netherlands and
Switzerland.
Lithobius muticus C . L. Koch
Lithobius muticus C. L. Koch, 1847: 151; Latzel, 1880b: 116; Brolemann, 1930:
317, figs 445-448; Verhoeff, 1936: 106; 1937: 210, fig. 58; Matic, 1966: 174, fig.
69 A-G.
Lithobius muticus is known from Austria, France, Germany, the Netherlands and
Switzerland.
Lithobius calcaratus C . L. Koch
Lithobius calcaratus C. L. Koch, 1844: 23, fig. 23; Latzel, 1880b: 105; Brolemann,
1930: 315, figs 442-444; Verhoeff, 1936: 105; 1937: 72.
Latzel was uncertain of the occurrence of L. calcaratus in Austria but it is known
from Belgium, France, Germany, the Netherlands, Scandinavia and Switzerland.
Lithobius lucifgus L. Koch
Lithobius lucijiugus L. Koch, 1862 : 82, fig. 38 ; Latzel, 1880b: 120;Brolemann, 1930:
320; Verhoeff, 1936: 106; 1937: 211, figs 29, 33-35; Matic, 1966: 178, fig. 71
A-F.
In this species the number of prosternal teeth is very variable and there is a
tendency for the regular rows of coxal pores to be supplemented by much smaller
pores (Verhoeff, 1937: fig. 29). Verhoeff (1935, 1937) used these characters as a
means of defining the subspecies latzeli, octodus and uarallemis but there is no
justification for this because the characters may vary among individuals from the
same breeding population or even from side to side of the same specimen, and if
these names are used at all they should have varietal status only. Individuals with
4 4 prosternal teeth are keyed separately.
Lithobius lucifgus is known from Austria, southern Germany, Switzerland and
Scandinavia and seems to be confined to relatively high altitudes.
+
Lithobius melamps Newport
Lithobius melamps Newport, 1845 (1844-45) : 371 ; Brolemann, 1930: 263, figs 374,
402, 403; Verhoeff, 1936: 104; 1937: 202; Matic, 1966: 136, fig. 52 A-D.
18
E. H.EASON
Lithobius glubratus C. L. Koch, 1847: 149; Latzel, 1880b: 74.
Lithobius melanops is known from Austria, France, Germany, the Netherlands,
Scandinavia and Switzerland.
Lithobius tricuspis Meinert
Lithobius tricuspis Meinert, 1872: 16; Latzel, 1880b: 76; Brolemann, 1930: 281, figs
423-426; Verhoeff, 1936: 104; 1937: 202.
Two characters usually found in this species, the coxolateral spines (VaC) on the
15th legs and the third pair ofspurs on the female gonopods, are sometimes absent
so that some specimens are difficult to identify. The form without 15 VaC is keyed
separately .
Lithobius tricuspis is known from ,4ustria, France, Germany, the Netherlands and
Switzerland.
Lithobius agilis C. L. Koch
Lithobius agilis C . L. Koch, 1847: 149; Latzel, 1880b: 78; Brolemann, 1930: 285,
figs 427-429; Verhoeff, 1936: 103; 1937: 202; Matic, 1966: 153, fig. 59 A.
Lithobius agilis pannonicus Loksa, 1948: 10, fig. 3 ; Matic, 1966: 153, fig. 59 B, C.
The shape of the spurs on the gonopods of the female neotype of L. agilis from
Bavaria (Eason, 1972: fig. 1 ) is much the same as that of the Hungarian specimens
which Loksa referred to the subspecies pannonzcus (Loksa, 1948: fig. 3), whereas
Brolemann figured much stouter spurs on females from the Pyrenees (Brolemann,
1930: fig. 429). Latzel described these spurs in agilis as long and slender and it is
likely that spurs of the shape figured by Loksa and also by Matic (1966: fig. 59 B,
C) are more typical of the northern form of the species than those figured by
Brolemann.
Lithobius agilis is known from every country in north-west Europe.
Lithobius erythrocephalus C. L. Koch
Lithobius erythrocephalus C . L. Koch, 1847: 150; Latzel, 1880b: 110, fig. 29;
Brolemann, 1930: 293; Verhoeff, 1936: 105; 1937: 214; Matic 1966: 157, fig. 61
A, B.
Brolemann recorded L. erythrocephalus from the Alps but not from northern or
central France. Authentic nort h-west European records of L . erythrocephalus are
from Austria, Germany, the Netherlands, Scandinavia and Switzerland, some of
those from Switzerland and most of those from Austria being referable to the
subspecies schuleri Verhoeff.
Lithobius erythrocephalus schuleri Verhoeff
Lithobius erythrocephalus: Latzel, 1880b: 110 (partim) ; Eason, 1964: 219, figs
397-404.
Lithobius erythrocepha1u.s schuleri Verhoeff, 1925: 144; 1937: 213, fig. 49.
Lithobius erythrocephalus schulleri: Matic, 1966: 159, fig. 61 C.
Among the characters given by Verhoeff (1925, 1937) for distinguishing schuleri
from the nominate subspecies of L . er_vthrocephalus are the absence of dorsal
NORTH-WEST EUROPEAN LITHOBIOMORPHA
19
flattening of the male 15th tibiae and the presence of small posterior projections on
T.13. But neither of these characters is reliable (Eason, 1972) and the two
subspecies can only be separated by the characters given in the key (p. 30),
namely the shape of the spurs and the size of the dorsolateral setae on the female
gonopods. The identity of the nominate subspecies has been established by the
designation of a female neotype from Bavaria (Eason, 1972): the shape of the spurs
on the gonopods of this specimen (Eason, 1972: fig. 2) and also of females of L.
erythrocephalus from Iceland (Eason, 1970b: fig. 4) is in marked contrast to the
shape of those ofschuleri (Verhoeff, 1937 : fig. 49). Latzel, who described these spurs
as “elongata et plerumque gracilia” in his diagnosis of L. erythrocephalus, probably
included both subspecies in his account of Austrian and Hungarian examples.
Verhoeff described schuleri originally from a number of mountainous localities in
Hungary and later recorded it from Switzerland where the typical form of the
species is also found (Verhoeff, 1935). Loksa (1955) mentioned both subspecies as
occurring in the Carpathians (Hungary) and Kaczmarek (1979) included them
both in her key to the Polish centipedes. Matic found schuleri to be widely
distributed in Rumania with two records of the typical form from the extreme
north of the country (Matic, 1966: fig. 62). The Austrian specimens used for a
description of L. erythrocephalus (Eason, 1964) all belong to schuleri and it has been
found in a number of Austrian localities (Eason, unpubl.) . All these records suggest
that the ranges of L. e. erythrocephalus and L. e. schuleri may overlap and, if the
difference between them is indeed clear-cut with no intermediate forms, they may
belong to two closely similar but distinct species.
Lithobius lusitanus Verhoeff
Lithobius borealis: Meinert, 1872: 322 (non Meinert, 1868); Latzel, 1880b: 90.
Lithobius lusitanus Verhoeff, 1925: 142, 148, fig. 2; 1937: 212.
Archilithobius stramineus Attems, 1927 : 245, figs 1-3, syn. nov.
Lithobius stramineus: Brolemann, 1930: 286; Serra, 1978: 36, figs 1-4.
Verhoeff s original description of this species, though brief, is easily
recognizable. Attem’s description of A . stramineus which was copied by Brolemann
was based on an unusually large specimen so that the synonymy of stramineus with
lusitanus has been overlooked. The species has been fully described by Serra under
L. stramineus. The typical form of L. lusitanus seems to be confined in Europe to the
Iberian peninsula, and in north-west Europe it is represented by the subspecies
valesiacus Verhoeff.
Lithobius lusitanus valesiacus Verhoeff
(Figs 8-12)
Lithobius lusitanus valesiacus Verhoeff, 1935: 196; 1937: 212, 224.
?Lithobius borealis: Verhoeff, 1936: 103 (non Meinert, 1868).
The subspecies valesiacus differs from the typical form of L. lusitanus in its smaller
size, reduced anterior spinulation of the legs and the complete absence of the spine
VaT. Males of valesiacus are often, like those of the typical form, without
modification of the posterior legs but large specimens may have a shallow but
distinct dorsal sulcus on the 15th tibia and are keyed separately.
Although the form appearing under L. borealis Meinert in Verhoeffs 1937 key
E. H. E.4SON
20
a
0
TO
I
I
0.2 mm
12
02mm
0.2 mm
0.2 mm
Figures 8- 12. Lithobiur lusilonus oalesiacus. Fig. 8, Right ocelli and organ ofTOmOsvary (TO);Figs 9, 10,
prosternum, ventral, showing variation in shape ofshoulder; Fig. 1 1 , E gonopods, ventral; Fig. 12, left
2 gonopod, lateral.
belongs to L. lapidicola Meinert, the L. borealis of his 1936 key is described as being
10-12 mm long (L.lapidicola rarely exceeds 9.5 mm in length) and seems more
likely to belong to L. lusitanus valesiacus. Many published records of L. borealis and
L. lapidicola from north-west Europe and elsewhere may also refer to valesiacus,
notably those from Mont Genkvre close to the Franco-Italian border mentioned by
Brolemann ( 1930) in his first remark supplementary to his main description of L.
NORTH-WEST EUROPEAN LITHOBIOMORPHA
21
lapidicola. The only previous authentic records of the subspecies, however, are the
original one from the neighbourhood of the Lake of Bienne (Verhoeff, 1935) and a
subsequent record from Stalden (Verhoeff, 1937), both in Switzerland.
The Swiss specimens described below add little to our knowledge of the
distribution of L. lusitanus ualesiacus but those from the Steigerwald constitute the
first German record.
Material examined
Seven specimens specimens from Switzerland collected by Dr Hans Jungen and
his students : four males from Suret between Aarau and Kupperswil ; three females
from Granichen.
Three specimens from the Steigerwald east of Wurzberg, Germany: a male and
a female collected by Mr H. C. Frund and a male collected by Mrs A. M. Albert.
Description
Size: 10-13mmlongand 1.3-1.7mm broadat T.lO. Head: 1.1-1.4mm broad;
broader than long, broader than T.3 but narrower than T.5; no median
thickening of posterior marginal ridge ; posterior border straight or very slightly
emarginate ; paired posterior depressions faint. Antennae : a third to two-fifths of
body length; of 27-38 (usually 34-36) articles, as broad as long or slightly
transverse, the last moderately elongate. Ocelli : 10-12 on each side; posterior
ocellus larger than posterosuperior; 1 +4,4, 3 ; 1 +4, 4, 2; 1 +4, 3, 3 or 1 +4, 3, 2
in straight horizontal rows (Fig. 8). Organ ofTomosvary: about the size of a small
ocellus (Fig. 8). Prosternum : with 2 2 teeth, the line of their apices distinctly
recurved ; porodont setiform, slightly stouter than a large seta; lateral to the
porodont the free border forms a broad shoulder, more marked in some specimens
than in others (Figs 9, 10) ; one specimen has an adventitious tooth on one side
lateral to the porodont and close to the point of the shoulder. Tergites: smooth
with little wrinkling; T. 1 narrower than T.3, approximately rectangular with
posterior border straight or very slightly emarginate; posterior borders of T.3 and
subsequent large tergites increasingly but never strongly emarginate; posterior
angles of T.8 evenly rounded, those of T.9 obtuse or squared, those of T.10
abruptly rounded, those of T. 1 1 with traces of projections or small projections,
those of T. 12 blunt, those of T. 13 with small or moderate projections, those of T. 14
blunt or angulated. Intermediate tergite : posterior border straight or slightly
emarginate. Coxal pores: 2, 4, 4, 3-4, 5, 5, 5 (frequently 3, 4, 4, 3) ; circular,
separated from one another by their own diameter or more. Tarsal articulations of
anterior legs: distinct. 14th and 15th legs: moderately thickened in both sexes, with
a shallow dorsal sulcus on the 15th tibia of larger males only; accessory apical claw
about half as long as principal claw on both legs; 15th about a third of body
length. Sensory spurs: as long as or longer than accessory apical claw on first to
13th legs, half as long on 14th, very short on 15th. Glandular pores: concentrated
on 14th and 15th legs with a few on 13th. Male genitalia: second genital sternite
without setae; gonopod of a single article with two or three setae. Female
gonopod: with two stout spurs and a claw with both medial and lateral denticles
well-developed (Fig. 1 1) ; one female has a third spur on one gonopod; dorsolateral
setae no stouter than general setae, usually three on the second article and one or
two on the third (Fig. 12); no dorsomedial setae.
+
E. H.EASON
22
Spiriulation :
C
1
2
3-4
5-6
7
8
9
10
11
12
13
14
t
Ventral
P
T
m
rn
m
m
m
m
m
m
m
rn
m
m
15
Letters in brackets indicate the usual variable spines. In addition to these
variations 1-4 VpP, 15 VaF, 4-6 VpF and 13 DaC are each present in one
specimen only; and 13 Vmt, 1 1 VpP, 2 VmF, 13 VpF, 1 DpP and 8 DpT are each
absent in one specimen only.
Lithobius borealis Meinert
Lithobius borealis Meinert, 1868: 263.
Lithobius lapidicola: Latzel, 1880b: 106; Brolemann, 1930: 291 ; Matic, 1966: 180,
fig. 70 &D. (non Meinert, 1872; non Verhoeff, 1937)
Lithobius lusitanus wiirmanus Verhoeff, 1937: 212, 224, fig. 54.
Lithobius saalachiensis Verhoeff, 1937: 214, 227, fig. 51; Jeekel, 1964: 134.
Non Litbbius borealis: Meinert, 1872: 322; Latzel, 1880b: 90.
Non Lithobius borealis: Verhoeff, 1936: 103.
Non Lithobius borealis: Verhoeff, 1937 : 2 1 1.
Non Lithobius borealis: Brolemann, 1930: 290.
Posterior projections on T. 1 1 and 13 are usually well-developed in examples of
L. borealis from the northern part of its range but may be inconspicuous in those
from further south.
Latzel’s description of L. borealis was copied from Meinert’s later (1872) account
of the species which was based on specimens of L. lusitanus (Eason, 1974). As
already suggested (p. 191, the L. borealis of Verhoeff s 1836 key probably belongs to
L. lusitanus valesiacus; and the L. borealis of Verhoeff s 1937 key certainly belongs to
L . lapiduola. Brolemann’s description of L. borealis is a copy of Meinert’s later
(1872) account combined with the spinulation of another species, probably L.
melamps (Eason, 1974).
Having been subjected to so much misdetermination it is difficult to be sure of
the distribution of L. borealis in north-west Europe, but there are authentic records
of the species, most of them under one or other of its synonyms, from Austria,
France, Germany and Scandinavia.
Lithobius lapidicola Meinert
Lithobius lapidicola Meinert, 1872: 328; Eason, 1980a: 26, figs 3, 4.
Lithobius pusillus Latzel, 1880b: 108; Brolemann, 1930: 294; Verhoeff, 1936: 105;
1937 : 216, syn. nov.
NORTH-WEST EUROPEAN LITHOBIOMORPHA
23
Lithobiuspusilluspusillzfater Verhoeff, 1925: 152; 1937 : 2 16; Brolemann, 1930: 295.
Lithobius borealis: Verhoeff, 1937 : 21 1 (non Meinert, 1868).
?Lithobius sulcatus L. Koch, 1862: 73, fig. 32.
Non Lithobius lapidicola: Latzel, 1880b: 106; Brolemann, 1930: 291 ; Matic, 1966:
180.
Non Lithobius erythrocephalus var. lapidicolus: Verhoeff, 1936: 105.
Non Lithobius lapidicola: Verhoeff, 1937 : 214.
Non Lithobius lapidicola: Jeekel, 1964: 135.
Lithobius pusillus as described by Latzel and subsequent authors is distinguished
from L. lapidicola by having only five or six ocelli on each side, no wrinkling of the
large tergites, no posterior projections on T.13 and by the absence of a ventral
spine on the 14th tibia. However, among the specimens in the Meinert Collection
in the Zoological Museum, Copenhagen, are two males and five females labelled
“Bona, Hispana”. Owing to Meinert’s method of labelling it cannot be said
whether these specimens come from BBne (Algeria) or from Spain but they
undoubtedly belong to L. lapidicola and have only minimal wrinkling of the large
tergites, no posterior projections or only traces of projections on T.13 and no
ventral spines on the 14th tibia : they differ from L. pusillus only in having seven or
more ocelli on each side. But Anderson (1980) found as few as six ocelli on each
side among the type specimens of L. lapidicola so that the number of ocelli is so
variable in this species that L. pusillus was almost certainly based on specimens of L.
lapidicola. The name pusillus was first used in the genus Lithobius, with some
uncertainty, by von Heyden (1837) for a species of Symphyla and subsequently by
Sseliwanoff (1880) for a species of Lithobius not found in north-west Europe, but it
has usually been regarded as attaching to L. pusillus Latzel which now becomes a
junior synonym of L. lapidicola.
The lectotype of L. sulcatus from Grutz, near Nuremberg, has fused anterior
tarsal articulations and is a first post-larval stadium of some small species of
Lithobius. It has been suggested (Eason, 1972) that this specimen must belong to
some species with fused tarsal articulations such as Lithobius microps (sensu Verhoeff,
non Meinert), but this species (see below) which appears to be common in Italy,
has never been recorded from north of the Alps and L. sulcatus is more likely to
belong to L. lapidicola in which the anterior tarsal articulations are rather faint in
adults and may appear to be fused in the first post-larval stadium.
Latzel’s, Brolemann’s and Matic’s accounts of L. lapidicola all refer to L. borealis.
The form Verhoeff ( 1936) keyed as L. erythrocephalusvar. lapidicolus Mein. is difficult
to place but is unlikely to belong to L. lapidicola. The form appearing under L.
lapidicola in Verhoeffs 1937 key is quite unlike L. lapidicola and may be a female of
L. mutabilis. The females described under L. lapidicola by Jeekel probably belong to
L. subtilis (Eason, 1980b).
The distribution of L. lapidicola in north-west Europe is even more difficult to
assess than that of L. borealis. Verhoeff (1937) stated that borealis ( = lapidicola) is
found in Switzerland, Tirol and northern Europe, and Anderson (1980) recorded
the species from several localities in Sweden. Most records of L. pusillus and L.
pusillus pusillifrater are from south of the Alps but Latzel described pusillus originally
from Austria, Rothenbuhler (1899) recorded it from Switzerland and Jeekel
(1977) from the Netherlands, so that L. lapidicola is probably quite widespread in
north-west Europe.
24
E. H. EASON
Lithobius microps Meinert
Lithobius mkrops Meinert, 1868: 265 ; Verhoeff, 1936: 105;Jeekel, 1964: 139.
Lithobius duboscqui Brolemann, 1896: 116; 1930: 328, figs 453-455.
Monotarsobius duboscqui: Verhoeff, 1937 : 189.
Lithobius (Monotarsobius) duboscqui: Matic, 1966: 222, fig. 89 A-D.
Non Lithobius microps: Meinert, 1872 : 330.
Non Lithobius microps: Brolemann, 1930: 322.
Non Monotarsobius microps: Verhoeff, 1937 : 189.
Verhoeff (1937) and many other authors, influenced by Meinert’s later (1872)
description of L. microps which is composite and was based on a number of small
species of Lithobius from Spain and North Africa in addition to L. microps (Eason,
1974), have attached the name microps to a Mediterranean species which may be
identical with L. biporus Silvestri, 1894 from Italy. However, in his 1936 key
Verhoeff seems to be referring to the true L. microps although he gave the number of
antennal articles as 26-29, whereas in north-west Europe this number is usually 25
and may be as low as 22 in examples of the species from north-west Germany
Ueekel, 1964). Brolemann (1930),on the other hand, attached the name microps to
a species from the Alpes-Maritimes which is quite different both from L. microps
and from L. biporus.
Lithobius microps is known from Belgium, France, Germany, the Netherlands,
Scandinavia and Switzerland, usually being synanthropic in the northern part of
its range.
Verhoeff (1937) included all the small species of Lithobius with fused anterior
tarsal articulations in the genus Monotarsobius Verhoeff, regardless of the number of
their antennal articles, thus including the last species. But many authors either
disregard Monotarsobius altogether or regard it as a subgenus of Lithobius. It is
considered here to be a subgenus of Lithobius but should include only those species
with the antennal articles limited to 20 or thereabouts as originally proposed by
Verhoeff ( 1905).
Lithobius (Monotarsobius) curtipes C. L. Koch
Lithobius curtipes C . L. Koch, 1847: 50; Latzel, 1880b: 130; Brolemann, 1930: 324.
Lithobius (Monotarsobius) curtipes: Verhoeff, 1936 : 106.
Monotarsobius curtipes: Verhoeff, 1937 : 188.
Lithobius (Monotarsobius) baloghi Loksa, 1947: 76, figs 10-16; Matic, 1966: 213, fig.
85 A-H.
Non Lithobius (Monotarsobius) curtipes: Matic, 1966: 2 16.
Loksa misdetermined Lithobius sseliixanoji Garbowski ( = L. pusillus Sseliwanoff)
as L . curtipes and described L. curtipes as a new species, L. baloghi Loksa. This
nomenclature was adopted by Matic.
Although Latzel did not record L. curtipes from Austria as defined by its present
boundaries it is known from France, Germany, the Netherlands, Scandinavia and
Switzerland.
Lithobius (Monotarsobius) crassipes L. Koch
Lithobius crassipes L. Koch, 1862 : 7 1, fig. 3 1 ; Latzel, 1880b: 128 ;Brolemann, 1930:
325, figs 449-452.
NORTH-WEST EUROPEAN LITHOBIOMORPHA
25
Lithobius (Monotarsobius) crassipes: Verhoeff, 1936 : 106; Matic, 1966: 206, fig. 82
A-D.
Monotarsobius crassipes: Verhoeff, 1937 : 188.
Lithobius crassipes is known from Austria, France, Germany, the Netherlands,
Scandinavia and Switzerland.
Lithobius (Monotarsobius) aeruginosus L. Koch
Lithobius aeruginosus L. Koch, 1862: 74, fig. 33; Latzel, 1880b: 126; Brolemann,
1930: 321.
Lithobius (Monotarsobius) aeruginosus: Verhoeff, 1936: 106; Matic, 1966: 199, fig. 78
A, B.
Monotarsobius aeruginosus: Verhoeff, 1937: 189, figs 15-17, 62, 63.
Lithobius aeruginosus is known from Austria, Germany, the Netherlands and
Switzerland.
Lithobius (Monotarsobius) austriacus (Verhoeff)
Monotarsobius austriacus Verhoeff, 1937 : 189, 192, fig. 65; Folkmanova, 1945: 126;
Borek, 1969: 33, figs 1-3.
Lithobius (Monotarsobius) austriacus: Loksa, 1947 : 84.
Verhoeff described this species from a single male from Turkenloch in Austria
which remains the only north-west European record. Loksa suggested that L.
austriacus might be no more than a variety of L. aeruginosus, but there have been a
number of records of the species, all from eastern Europe, and both Folkmanova
and Borek have supplemented Verhoeff s rather scanty original description.
Family Henicopidae
Genus Lamyctes Meinert
Lamyctes fulvicornis Meinert
Lamyctes fulvicornis Meinert, 1868: 267 ; Brolemann, 1930: 333, figs 456-458 ;
Verhoeff, 1936: 106; Matic, 1966: 241, fig. 98 A-E.
Henicops fulvicornis: Latzel, 1880b: 133, figs 31-34.
Only females of Lamyctes fulvicornis are found in north-west Europe, where it is
known from Austria, France, Germany, the Netherlands and Scandinavia.
Genus Lamyctinus Silvestri
Lamyctinus coeculus (Brolemann)
Lithobius coeculus Brolemann, 1889: 273, pl. 1, fig. 1.
Lamyctinus coeculus: Silvestri, 1909: 39, fig. 1, 1-1 1.
Lamyctes coeculus: Brolemann, 1930; 336, figs 463-465.
The genus Lamyctinus, which Silvestri (1909) based on the absence of ocelli, was
disregarded by Brolemann (1930).But Anderson (1970b) has shown a significant
difference in larval development between coeculus and Lamyctes fulvicornis and
believes that the genus Lamyctinus should be retained.
Lamyctinus coeculus is known from natural habitats in the Ethiopian and
neotropical regions but is exclusively synanthropic in north-west Europe where
females have been found in greenhouses in Paris (Brolemann, 1930), Copenhagen
and Goteborg (Enghoff, 1975).
E. H . EASON
26
KEY TO THE NORTH-WEST EUROPEAV LITHOBIOMORPHA
Most of the terms and notations used in the following key are in common use but
some of those used to describe the male and female genitalia require explanation.
The second genital sternite in the male (the small ventral plate between the
gonopods) is abbreviated to ‘G.S.2’.
Of the modified setae found on the dorsum of the female gonopod, those on the
basal article just proximal to the insertions of the spurs (Fig. 7) are referred to
simply as the ‘dorsomedial setae’ : and those arranged along the intercondylar line
of the second article (Figs 7, 12) are referred to as the ‘dorsolateral setae’. The setae
found elsewhere on the gonopod are referred to as the ‘general setae’.
Most of the characters used in the key can be seen quite easily without
dissection. But, although the dorsolateral setae of the female gonopod can usually
be seen in the intact specimen, it is necessary to remove the gonopod in order to see
the dorsomedial setae.
1.
-
Coxal pores numerous and arranged irregularly
.
Coxal pores less numerous and arranged in a single row on each
coxa .
2.
-
3.
-
4.
4
5
A single ocellus on each side. Length 7-1 1 mm Lamyctes fuluicornis
No ocelli. Length 3.5-5mm .
,
Lamyctinus coecutus
5. Prosternal teeth small or vestigial. Porodont usually massive
Prosternal teeth well-developed. Porodont usually slender
-
3
Posterior projections on T.6 prominent and angulated.
General setae on 15th legs very short. A single row of ventral
seriate setae on distal half of 15th metatarsus. Glandular
pores of 15th legs numerous and dense, on all aspects of
femur, tibia, tarsus and metatarsus and also on medial aspect
of prefemur. 15th accessory apical claw present
. Eupolybothrus grossipes
Posterior projections on T.6 feeble and rounded or absent.
General setae on 15th legs fairly long. No ventral seriate setae
on 15th metatarsus. Glandular pores of 15th legs sparse, on
medial aspects of femur, tibia, tarsus and metatarsus only.
15th accessory apical claw absent .
. Eupolybothrus tridentinus
Marginal ridge of head not interrupted laterally. No true spines on
any leg but a sharp distal anterior projection on first to 1 lth tibiae
in the position of DaT. An anterior and a posterior 15th accessory
apical claw .
Marginal ridge of head interrupted and slightly projecting
laterally. Always with spines on some legs but no tibia1 projections.
With or without a single (posterior) 15th accessory apical claw
-
2
.
.
6
7
6. T.9, 1 1 and 13 with posterior projections. First legs
thickened, without spines. 15 VaC absent. 15th accessory
apical claw present. Male 15th tibia with a dorsal sulcus
.
Harpolithobius anodus
NORTH-WEST EUROPEAN LITHOBIOMORPHA
-
27
T.9. 11 and 13 without posterior projections. First legs not
thickened, with spines. 15 VaC present. 15th accessory apical
. Lithobius castaneus
claw absent. Male 15th legs not modified
7. A ventral medial spine (VmC) present on 15th coxa
. Lithobius pilicornis
-
15 VmC absent
.
8
8. 4 + 4 or more prosternal teeth
2 2 or 3 3 prosternal teeth
-
+
+
.
.
9
12
9. Tergites without posterior projections or with feeble projections on
T.13 only .
. Lithobius lucijiugus
Three or more of the last short tergites with posterior projections 10
10. T.6, 7, 9, 11 and 13 with posterior projections. T.l much
broader than T.3. Female gonopod with numerous
. Lithobius validus
dorsomedial setae
Only T.9, 11 and 13 with posterior projections. T.l not
broader than T.3. Female gonopod without dorsomedial
setae .
11
11. Usually 4 + 4 prosternal teeth. 15 VaC and 15th accessory apical
.
.
Lithobius piceus
claw present. Male G.S.2 without setae
- 5 + 5 or more prosternal teeth. 15 VaC and 15th accessory apical
. Lithobius forjicatus
claw absent. Male G.S.2 with two setae
12. Three to six ocelli on each side, in a single horizontal row.
Tarsal articulations of first to 13th legs fused .
Five or more ocelli on each side, in an irregular mass or in
two or more horizontal rows. Tarsal articulations of first to
13th legs fused or not
.
13
15
13. Antennae usually of 25 articles. VaF, VaT, DaF and DpF
altogether absent. 15th accessory apical claw present Lithobius microps
- Antennae usually of 20 articles, rarely more than 21. VaF, VaT,
DaF and DpF present on some legs. 15th accessory apical claw
absent .
14
14. In males 15th prefemur with VaP, VpP, DmP and DpP
reduced in size, at most only twice as long as broad, with VpP
in the form of a small trifid spur; 15th femur barely twice as
long as broad; 14th legs similarly but less markedly modified
. Lithobius austriacus
In males 14th and 15th legs not modified apart from simple
thickening; 15th prefemoral spines about five times longer
than broad; 15th femur three times longer than broad
.
Lithobius aeruginosus
15. Antennae usually of 20 articles, rarely more than 21. Tarsal
articulations of first to 13th legs fused .
Antennae of 28 or more articles. At least a trace of tarsal
articulations on first to 13th legs .
16
17
E. H . EASON
2s
~
16. Posterior ocellus larger than adjacent ocellus. DmP usually
present on first leg, always on fifth and always extending
further forward than DaP. Male 15th tibia thickened without
projection. Male G.S.2 with two setae .
. Lithobius crassipes
Posterior ocellus smaller than adjacent ocellus. DmP never
present further forward than 1 lth leg and rarely extending
further forward than DaP. Male 15th tibia thickened with a
distal dorsal dorsal blunt projection. Male G.S.2 without
setae .
. LithobiuJ curtipes
17. T.9, 11 and 13 with posterior projections
.
Tergites without projections or with small projections on T. 11 and
13 or T.13 only .
18
-
18.
-
15th accessory apical claw absent .
15th accessory apical claw present.
28
19
20
19. 15 VaF absent. DmP starts on 7-10. Male 15th tibia with a
. Lithobius nodulipes
prominent dorsal subdistal setose node .
__
15 VaF present. DmP starts on 1-4. Male 15th tibia sometimes
with a dorsal sulcus but never with a node .
. Lithobius tenebrosus
20.
-
15 VaC present
15 VaC absent
.
21
.
23
2 1. Large tergites distinctly wrinkled. Spurs of female gonopod long
-
and slender, often expanded distally, rarely more than two. Claw
.
Lithobius agilis
of female gonopod distinctly tridentate.
Large tergites smooth or indistinctly wrinkled. Spurs of female
gonopod stout, or if long and slender never expanded distally,
usually more than two. Claw of female gonopod usually simple or
hidentate, rarely tridentate .
22
22.
+
3 3 prosternal teeth. Usually five or more pores on each of
the 13th and 14th coxae. 15th legs slender. Length 13-21 mm
.
Lithobius piceus
2 2, rarely 2 3 or 3 3, prosternal teeth. Rarely more than
four pores on each of the 13th and 14th coxae. 15th legs
slightly thickened. Length 10-16mm
.
. Lithobius tricuspis
+
+
+
23. Medial pair of prosternal teeth usually larger and often projecting
further forward than lateral pair. Male 15th femur and tibia each
with a dorsal sulcus
.
. Lithobius macilentus
Lateral pair of prosternal teeth projecting further forward than
medial pair, or both pairs on same level. Male 15th femur without
a dorsal sulcus
.
24
24. Male 15th, 14th and usually 13th tibiae each with a dorsal
sulcus or pit. Female gonopod with several dorsomedial setae
25
-
Male posterior legs not modified. Female gonopod without
dorsomedial setae .
.
27
NORTH-WEST EUROPEAN LITHOBIOMORPHA
25.
-
29
14-23 ocelli on each side. Antennae usually of more than 44
articles. Free margin of prosternum extending straight laterally
beyond porodonts and then sloping quite sharply backward.
Posterior projections on T.9 well-developed. Length 11-18 mm
. Lithobius dentatus
6-16 ocelli on each side. Antennae rarely ofmore than 44 articles.
Free margin of prosternum sloping obliquely backward
immediately lateral to porodont. Posterior projections on T.9
usually feeble. Length 6-1 1.5mm .
26
26. 9-16 ocelli on each side, posterior ocellus the largest.
Porodont well-separated from lateral prosternal tooth. In
males 14th tibia with a feeble dorsal sulcus on its distal half
only, 15th tibia with an inconspicuous dorsal subdistal pit.
Male G.S.2 without setae. Length 7.5-1 1.5 mm
. Lithobius subtilis
Six to nine ocelli on each side, posterosuperior ocellus usually
the largest. Porodont immediately adjacent to lateral
prosternal tooth. Male 14th and 15th tibiae each with a
broad deep dorsal sulcus. Male G.S.2 with two setae. Length
6-8 mm
. L ithobius pygmaeus
27. Free margin of prosternum with broad irregular shoulders lateral
to porodonts. Female gonopod with two, rarely three, short stout
spurs and a distinctly tridentate claw .
. Lithobius melanops
Free margin of prosternum sloping obliquely backward
immediately lateral to porodonts. Female gonopod with three,
rarely two, fairly long spurs and the claw simple or bidentate,
rarely tridentate .
. Lithobius tricuspis
28. 15th accessory apical claw absent .
15th accessory apical claw present.
29
34
29. First three or four large tergites each with a pair of curved
paramedian sulci arising from the posterior marginal sulcus. 15
VaC present
.
. Lithobius castaneus
___
Large tergites without sulci. 15 VaC absent .
30
30. When a single ventral spine is present on the 15th tibia it is
usually VmT. Male posterior legs not modified Lithobius lucijiugus
- When a single ventral spine is present on the 15th tibia it is
usually VaT. Male 14th and/or 15th legs modified
.
31
Head as broad as T.5 or broader. In males 14th tibia with a feeble
setose swelling immediately proximal to DpT, 15th leg not
modified. Male G.S.2 with two setae .
. Lithobius muticus
Head as broad as T.3. Male 15th leg modified. Male G.S.2
.
32
without setae
32. In males 14th and 15th tibiae each with a dorsal sulcus
associated with groups or lines of setae, 15th femur without a
sulcus. Length 8-1 1 mm
.
.Lithobius latro
E. H.EASON
30
-
In males 14th leg not modified, 15th femur and tibia each
with a dorsal sulcus. Length 10-16mm .
33
33. Male 15th prefemur with a marked distomedial projection. Male
intermediate tergite without a central depression . Lithobius cyrtopus
Male 15th prefemur slightly protuberant distomedially but
without a marked projection. Male intermediate tergite with a
. Lithobius pelidnus
central depression
34.
-
15 VaC present. Spurs of female gonopod not less than three
times longer than broad, often slightly expanded distally
15 VaC absent. Spurs offemale gonopod not more than three
times longer than broad, not expanded distally
.
35
36
35. Female gonopod with spurs about three times longer than broad
and dorsolateral setae as stout as or stouter than general setae
. Lithobius erythrocephalus erythrocephalus
Female gonopod with spurs four or five times longer than broad
and dorsolateral setae more slender than general setae
.
Lithobius erythrocephalus schuleri
36. Male 15th femur modified .
Male 15th femur not modified
-
37
.
38
37. Posterior ocellus relatively small, preceded by a larger
intermediate ocellus and those of the main mass in a rosette. In
males 15th femur with a distomedial wart-like outgrowth but no
sulcus, neither 15th prefemur nor tibia modified. Claw of female
gonopod bidentate
.
. Lithobius calcara tus
Posterior ocellus large, preceded by those of the main mass in
curved horizontal rows. In males 15th femur with a dorsal sulcus
and sometimes a feeble distomedial setose swelling but no wart-like
outgrowth, 15th prefemur with a distomedial projection and 15th
tibia with a dorsal sulcus. Claw of female gonopod tridentate
. Lithobius cyrtopus
38.
-
In males 15th and usually 14th tibia with a dorsal sulcus or
pit
.
Male posterior legs not modified .
.
39
42
39. Lateral pair of prosternal teeth projecting distinctly further
-
forward than medial pair. In males 14th tibia without a sulcus,
15th tibia with a very feeble dorsal sulcus. Female gonopod with
dorsolateral setae no stouter than general setae, no dorsomedial
setae .
. Lithobius lusitanus valesiacus
Both pairs of prosternal teeth on much the same level. Male 14th
tibia with a dorsal sulcus. Female gonopod with the more distal
dorsolateral setae stouter than general setae, several small
40
dorsomedial setae .
40.
In males 15th tibia with an inconspicuous dorsal subdistal
pit, 14th and usually 13th tibia with a feeble dorsal sulcus.
Male G.S.2 without setae. The two distal dorsolateral setae of
NORTH-WEST EUROPEAN LITHOBIOMORPHA
-
41.
-
31
female gonopod conspicuously longer and stouter than the
. Lithobiussubtilis
other dorsolateral setae .
In males 15th, 14th and usually 13th tibiae each with a
distinct dorsal sulcus. Male G.S.2 with two setae. The two
distal dorsolateral setae of female gonopod not conspicuously
41
longer and stouter than the others
13-20 ocelli on each side, posterior ocellus the largest. Tibial sulci
in males associated with groups or lines of setae. Length 10-15 mm
. Lithobius mutabilis
6-9 ocelli on each side, posterosuperior ocellus usually the largest.
Tibial sulci in males not associated with groups or lines of setae.
Length 6 to 8mm
.
. Lithobius pygmaeus
42.
-
15th prefemur in both sexes with a supplementary medial
spine between DpP and VpP
.
. Lithobius borealis
15th prefemur without a supplementary spine
.
43
43. Free margin of prosternum with broad irregular shoulders lateral
to porodonts. Large tergites smooth or indistinctly wrinkled. 15
DaP always present. Length 10-1 3 mm
Lithobius lusitanus valesiacus
- Free margin of prosternum sloping obliquely backward
immediately lateral to porodonts. Large tergites often distinctly
wrinkled. 15 DaP usually absent. Length 6-9.5 mm
. Lithobius lapidicola
ACKNOWLEDGEMENTS
I wish to thank Dr Henrik Enghoff of Copenhagen University for enabling me to
examine the specimens of Lithobiidae in the Zoological Museum, Copenhagen, on
which a large part of this study is based; Mrs A. M. Albert ofGottingen University
and Mr H. C. Frund of Wurzburg University for lending me their specimens from
the Steigerwald; and Dr Hans Jungen of Zurich University for lending me
specimens of Lithobius pygmaeus and L. lusitanus valesiacus from Switzerland. My
thanks are also due to Mr Ernst Schatzmann of Zurich University for his accurate
observations on the Swiss specimens.
REFERENCES
ANDERSON, G., 1979a. Taxonomical Studies on the Post-embTonic Development in Lithobius, with a Brief Comparison
with Lamyctes (Chilopoda: Lithobiomorpha) . Dissertation, University of Goteborg.
ANDERSON, G., 1979b. On the use of larval characters in the classification of lithobiomorph centipedes
(Chilopoda, Lithobiomorpha). In M. Camatini (Ed.), Myriapod Biology: 73-81. London: Academic Press.
ANDERSON, G., 1980. Lithobius borealis Meinert and L. lapidicola Meinert in Sweden (Chilopoda: Lithobiidae).
Entomologica Scandinavica, 11: 45-48.
ATTEMS, C. G., 1927. Myriopoden aus dem nordlichen und ostlichen Spanien, gesammelt von Dr F. Haas in
den Jahren 1914-1919. Abhandlungen hrsg. von der Senckenbergischen Natulforschenden Gesellschaft, 39: 233-290.
BOREK, V., 1969. NBlez Gynandromorfa Monotarsobius austriacus Verhoeff, 1937 (Chilopoda), Acfa Musei
Reginaehradecensis S.A: Scientiae Naturales, 10: 33-34.
BROLEMANN, H. W., 1889. Contributions i la Faune Myriadopologique MCditerrantenne. Trois espkces
nouvelles. Annales de la Socie'te' Linnbnne de Lyon, 1888 (N.S.), 35: 271-282.
BROLEMANN, H. W., 1896. MatCriaux pour servir i une Fame des Myriapodes de France. Feuille des Jtrtnrr
Naturalistes, 26; 115-1 19.
E. H. EASON
32
BROLEMANN, H. W.,1898. Myriapodes d’Ahusquay (Basses Pyrinies). Feuille des Jeunes Naturalistes, 28:
200 -203.
BROLEMANN. H. W.,1930. % n e n t s d’une Faune des Myriapodes de France. Chilopodes. Faune de France, 25:
1-405.
DEMANGE, J.-M., 1958. Sur quelques Myriapodes cavernicoles de France et de Suisse. Revue Suisse de Zoologic,
6.5: 843-855.
EASON, E. H., 1964. Centipedes of the British Isles. London: Warne.
EASON, E. H.. 1970a. A redescription of the species of Eupolybothrus Verhoeff s. str. preserved in the British
Museum (Natural History) and the Hope Department of Zoology, Oxford (Chilopoda, Lithobiomorpha).
Bulletin of the British M u m (Xatural H k t o y l , <oology, 19: 289-310.
EASON, E. H., 1970b. The Chilopoda and Diploda of Iceland. Entmnologica Scandinavica, I: 47-54.
EASON, E. H., 1972. The type specimens and identity of the species described in the genus Lithobius by C. L.
Koch and L. Koch from 1841 to 1878 (Chilopoda: Lithobiornorpha). Bulletin ofthe British Museum (.Vatural
,
105-150.
Hisbry), ~ o o l o g y 22:
EASON, E. H., 1974. The type specimens and identity of the species described in the genus Lithobius by F.
Meinert, and now presemed in the Zoological Museum Copenhagen University (Chilopoda:
Lithobiomorpha). <oological Journal of fhe Linnean Society,55: 1-52.
EASON, E. H., 1980a. On Lithobiidae from Sardinia (Chilopcda: Lithobiomorpha). Bulletin, <oologisch Museum,
Cniversiteit van Amstmdam. 7: 21-31.
EASOS, E. H., 1980b. On Lithobius subtifis Latzel, 1880, a little-known European species of Lithobiidae
[Chilopoda: Lithobiornorphai. Bulletin, <wlogisch Museum, Universiteit van .4msterdam, 7: 125- 131.
EASON, E. H. & Minelli, A., 1976. The identity of the species of Lithobiidae described by F. Fanzago and G.
Fedrizzi from 1874 to 1881 (Chilopoda, Lithobiomorpha). Fragment0 Entomologica, 12: 183-205.
ENGHOFF, H., 1975. Notes on Lamyctes roeculus (Brolemann), a cosmopolitic, parthenogenetic centipede
(Chilopoda: Henicopidaei. &fomologua Scandinavica, 6: 45-46.
FANZAGO, F., 1874. I chilopodi italiani. Monografia. .Ifti della Societa T‘eneto-Trentinadi Scienze .vaturak in Padova,
3: 17-64.
FEDRIZZI, G., 1877. I litobi italiani. Atti della Societd Venefo-Trentina di Sciewe .Vafurali in Padova, 5: 184 -233.
FOLKMANOVA, B., 1945. 0 ntkteech zajimav9ch stonoikach ze ziitopoveho bzemi berounske piehrady.
Entomologick4 Listy, 8: 12(t 129.
HAASE, E., 1880. Schlesiens Chilopoden, I. Chilopoda a m o r p h a . Inaugural Dissertation, University of Breslau.
HEYDEN, C. H. G. von, 1837. Lithobius? puillw in Kurzere Mittheilungen. Museum Senckenbergianum.
Abhandlungen aus dem Gebiete der beschreibenden .Ihturgeschichte, 2: 305.
JEEKEL. C. ‘4. W., 1964. Beitrag zur Kenntnis der Systematik und Okologie der Hundertfiisser (Chilopoda)
Nordwestdeutschlands. .4bhandlungen und Ik.handlungen des .2htunuissenschaftlichen Vereins in Hamburg, N. F., 8:
I 11-153.
JEEKEL. C. A. W., 1977. Voorlopige atlas van de verspreiding de Nederlandse duizendpoten (Chilopoda).
I’erslagen en Technkche gegeuens Inshtuut L ~ O OTaxonomkche
~
<oi?logi (<ovlogisch Museum), uniuersiteit van Amsterdam,
13: 1-55.
KACZMAREK, J., 1979. Pareczniki (Chilopoda) Polski. Poznan: Drukarnia Uniwenytetu im. Adama
Mickiewicza.
KOCH, C. L., 1844. Deutschlands Crustaceen, :Myriapoden und Arachniden, 40. Regensburg.
KOCH, C. L., 1847. System der Myriapcden. In G. Panzer & A. Herrich-Schaffer (Eds), Kritisch Revision der
Insectenfauna Deutschlands, 3. Regensburg.
KOCH, L., 1862. Die Myriapodengatfung Lithobius. Numberg: J . L. Lotzbeck.
LATZEL, R., 1880a. Zwei neue rnitteleuropaische Arten der Gattung Lifhobiw Leach. Zoologischer Anzezger, 3:
225-226.
LATZEL, R., 1880b. Die hljviopodm der 6sterreuhisch-L~ngarischen Monarchie, I . Die Chilopoden. Wien: Holder.
LINNE, C., 1758. $ys.!ema .\aturae, I. 10th ed. Holmiae.
LOKSA, I., 1947.Beitrage zur Kenntnis der Steinlaufer--,Lithobiiden-Fauna des Karpatenbeckens, 1. Fragment0
Faunistica Hungarica, 10: 73-85.
LOKSA, I., 1948. Beitrage zur Kenntnis der Steinlaufer-, Lithobiiden-Fauna des Karpatenbeckens, 2. Fragment0
Faunirhca Hungarica. 11: 1-1 I .
LOKSA, I., 1955. Uber der Lithobiiden des Faunagebiets des Karpatenbeckens. Acta <oologzca Academiae
Scientinrum Hungaricae, I : 33 1-349.
MATIC, Z., 1966. Clasa Chilopoda. Subclasa Anamorpha. Fauna Republicii Socialkfe Romtinia, 6: 1-272.
MEINERT, F.. 1868. Danmarks Scolopender og Lithobier. .Vaturhistorkk Tidsskrifl, 5: 241-268.
MEINERT, F., 1872. Myriapoda Musaei Havniensis: bidrag ti1 myriapodernes morphologi og systematik; I1
Lithobiini. .Vaturhktorisk Tidrskrlft, 8: 281-344.
NEWPORT, G., 1844. A list of the species of Myriapcda, order Chilopoda, contained in the cabinets of the
British Museum, with a synoptic description of forty-seven new species. Annals and Magazine of .Natural History,
13: 94-101.
NEWPORT, G., 184645. Monograph of the class Myriapoda, order Chilopoda; with observations on the
general arrangement of the Aniculata. Transactionr of the Linman Society ofLondon, 19: 265-302 (1844), 349--439
( 1845;.
NORTH-WEST EUROPEAN LITHOBIOMORPHA
33
ROTHENBUHLER, H., 1899. Ein Beitrag zur Kenntnis der Myriapodenfauna der Schweiz. Revue Suiue de
zoologic, 6: 199-271.
ROTHENBUHLER, H., 1901. Myriapoden Graubundens besonders des Engadins und des Munsterthales. Revue
Suisse de zoologic, 9: 357-377.
SERRA, A,, 1978. Contribucion a1 conocimiento de 10s Quilopodos de algunas zonas aridas de la Peninsula
Iberica. Los Monegros (Arag6n). Miscelinea zoologica, 4 ( 2 ): 31-42.
SILVESTRI, F., 1909. Contribuzioni alla conoscenza dei Chilopodi IIILIV. Bollettino del Laboratorio di zoologia
Generale e Agraria della R. Scuola Superiore d'Agricultura in Portici, 4 : 38 -65.
SSELIWANOFF, A., 1880. Materiali k izucheniyu russkikh tisyacbenogikh (Myriapoda). TruG Russkago
Entomologicheskago Obshchestva, 11: 3-26.
VERHOEFF, K. W., 1905. Uber die Entwicklungsstufender Steinlaufer, Lithobiiden, und Beitrage zur Kenntnis
der Chilopoden. <oologische Juhrbiicher (Supplement),8: 195-298.
VERHOEFF, K. W., 1925. Beitrage zur Kenntnis der Steinlaufer, Lithobiiden. Archiv f a r Naturgeschichte, 91:
124- 158.
VERHOEFF, K. W., 1935. Quer durch Schwarzwald und schweizerischen Jura (Chiemgau), Chilopoden.
Verhandlungen des ~atuwissenschaftluhenVereins in Karlsruhe, 31: 181-208.
VERHOEFF, K. W., 1936. Diplopoda, Symphyla, Pauropoda, Chilopoda. Die Tierwe16 Mit&leuropas, 2 ( 3 ):
1.- 120.
VERHOEFF, K. W., 1937. Chilopoden-Studien. Zur Kenntnis der Lithobiiden. Archivfar "Waturgeschichte,(N. F.)
6: 171-257.
2