A redescription of the species of Eupolybothrus Verhoeff s. str. preserved in the British Museum (Natural History) and the Hope departement of Zoology Oxford (Chilopoda Lithobiomorpha)
Author
E. H. Eason
text
Bulletin of the British Museum (Natural History), Zoology
1970
19
289
310
http://un.availab.le
journal article
Eason-1970-Eupolubothrus-E-grossipes
Eupolybothrus
(E.) grossipes (
C. L. Koch
)
Figs. 4 to 7
Lithobius grossipes
C. L. Koch, 1847
, p. 146; L. Koch, 1862, p. 32, fig. 4.
Lithobius montanus
C. L. Koch, 1847
, p. 148; L. Koch, 1862, p. 27, fig. 1.
Lithobius festivus
L. Koch, 1862
, p. 29, fig. 2.
Lithobius (Eulithobius) fasciatus
: Pocock, 1890, p. 61.
MATERIAL EXAMINED.
The following specimens, preserved in spirit, are present in the Koch Collection of
Arachnida
and
Myriapoda
in the British Museum (Natural History); except where otherwise stated the labels appear to be in L.
Koch's
hand:
"
Lithobius grossipes
C. Koch
,
festivus
L. Koch
"
"
Idria
"
(Reg. no.
13.6.18.262
).
This specimen,
a female
30 mm
long, although preserved in spirit has obviously been dried. An unusual feature is that the 14th leg appears stouter than the 15th; this appearance was noted, not only by L. Koch (1862) who undoubtedly used this specimen for his description of
L. grossipes
, but also by C. L. Koch (1847) in his original description. I therefore believe it to be the
holotype
and have labelled it accordingly. The fact that C. L. Koch gives
Trieste
as the
type
locality is consistent with the same specimen having been used for both descriptions since
Idria
(
Idrija
,
Yugoslavia
) is in the neighbourhood of
Trieste
.
"
Lithobius grossipes
C. K.
" "
Hopfgarten
,
26 August, 1868
" (Reg. no.
13.6.18.263-265
)
Three females
28 to 30 mm
long.
"
Lithobius grossipes
C.K.
" "
Seiseralpe
[an alpine hut in
Italy
], *[leg.] Gredler" (Reg. no.
13.6.18.266
).
This specimen,
a male
35 mm
long, agrees in detail with L.
Koch's
(1862) description of
L. montanus
C. L. Koch
, a description based on a single specimen sent him
by
Prof. P. Gredler
from
"
Seiseralpe
"
in
South Tyrol.
This is undoubtedly the specimen in question and Was presumably relabelled by L. Koch when he realised its identity with
L. grossipes
, of which
L. montanus
is now an accepted synonym.
"
grossipes
,
Eisarkthal
[
Valle
dell'Isarco
,
Italy
]
1870
" (Reg. no.
13.6.18.267-269
).
Two females and
a male
30 to 31 mm
long.
"
Lithobius grossipes
C.K.
,
festivus
L.K.
"
"
Sudtirol
"
(Reg. no.
13.6.18.270-282
, part).
Two females
35
and
37 mm
long, a mature male
34 mm
long and
two immature females
17
and
21 mm
long.
"
grossipes
,
Sudtirol
" (Reg. no.
13.6.18.270-282
, part).
A single female
37 mm
long.
"
Lithobius grossipes
C.K.
,
festivus
L.K.
"
"
Sudtirol
"
(Reg. no.
13.6.18.270-282
, part).
Four mature females
28 to 31 mm
long and
three immature females
16 to 21 mm
long.
"
Lithobius grossipes
C.K.
,
festivus
L.K.
"
"
Sudtirol
"
(Reg. no.
13.6.18.270-282
, part).
Three mature males
28 mm
long and
two immature males
12
and
14 mm
long.
"
Lithobius grossipes
C. Koch.
" "
Meran
[
Merano
,
Italy
], [leg.]
Milde
" (Reg. no.
13.6.18.283-292
, part).
Three females
27 to 30 mm
long, six mature males
27 to 38 mm
long and an immature male
19 mm
long.
"
grossipes
,
Meran
[
Merano
,
Italy
], [leg.]
Ausserer
" (Reg. no.
13.6.18.283-292
, part).
An immature male
23 mm
long.
"
Lithobius grossipes
C.K.
"
"
Garmisch
"
(Reg. no.
136.18.293
).
An immature male
24 mm
long. Although not agreeing in every detail with L.
Koch's
(1862) original description of
L. festivus
from
Garmisch
(
Bavaria
) this may be one of the specimens on which it was based.
"
grossipes
,
Ballino
[
Italy
]
12.IX.69
" (Reg. no.
13.6.18.298
).
An immature female
17 mm
long. The identity of this specimen is doubtful; it may possibly belong to
E. fasciatus
.
"
L. grossipes
." "
Susa
,
Oberitalien
" (Reg. no.
13.6.18.299-300
).
Two immature males
18
and
24 mm
long. The locality label is in L.
Koch's
hand but the identity label has been rewritten.
"
leptopus
?
Hopfgarten
" (Reg. no.
13.6.18.365
).
An immature female
13 mm
long.
The following specimens are in the British Museum (Natural History), preserved in spirit in a jar labelled "
Lithobius fasciatus
Newp.
Liguria
,
O. Thomas
" in
Pocock's
hand (Reg. no.
89.3.8.1-2
):
A total of
31 specimens
ranging in size from fourth larval stadia
9 mm
long to a mature male
43 mm
long. These specimens almost certainly come from
Busalla
(
Liguria
,
Italy
) both because Pocock (1890) mentions
Busalla
as the locality and
Mr. Oldfield Thomas
as the collector of examples of "
L. fasciatus
" in his paper on
Ligurian
centipedes, and because they were mixed with two specimens of
Lithobius doriae
Pocock
; in the same paper this author describes
L. doriae
as a
new species
not only from the same locality (
Busalla
) but also from the same habitat as "
L. fasciatus
" (Pocock, 1890: 59 & 64) and he must mistakenly have included specimens of the two species under the same label. I have relabelled these two examples of
L. doriae
and placed them in a separate tube.
The
following specimens were contained in the above jar along with the
Ligurian
specimens but in separate tubes:
A badly mutilated "stadium agenitalis"
12 mm
long labelled "
Eulith. grossipes
,
Grande Chartreuse
[
Isere
,
France
], [leg.]
A. Dollfus
" (Reg. no.
89.3.10.1
, part) and a badly mutilated immature male
15 mm
long labelled "
L.
grossipes
Koch
,
Portofino
[
Italy
] " (Reg. no.
89.3.10.1
, part). Both labels appear to be in
Pocock's
hand; in neither case is it possible to identify the specimens other than as species of the subgenus
Eupolybothrus
.
DIAGNOSIS or ADULT. Length
27 to 43 mm
. Antennae of 40 to 58 articles. Glandular pores of 15th leg concentrated on internal and dorsal aspects of prefemur and all aspects of other articles. 15th metatarsal general setae rarely exceeding a quarter the diameter of the article in length. 15th metatarsal seriate setae present. Basal pit of male 15th femur small and shallow. Internal dorsal sulcus of male 15th femur extending to margin of pore-free area which is not swollen. No coxolateral spines.
DESCRIPTION or ADULT. Length:
27 to 43 mm
; 15th legs up to two-thirds of body-length. Colour: pale to dark brown, often with a darker dorsal median band. Antennae: half to two-thirds of body-length, sometimes shorter in female; of 40 to 50 (female) and 47 to 58 (male) articles, the distal 10 to 14 often very elongate in large specimens. Ocelli: number and general pattern as described for
E. fasciatus
; in some specimens, however, the ocelli of the superior row may be round rather than oval and little larger than the others, but they are always relatively widely separated from one another. There is a tendency for the ocelli of the second, third and fourth rows to be more numerous than those of the superior row but this is by no means invariable. Prosternum: with 6+7 to 8+9 small teeth, usually 7+7 or 8+8; minute setiform lateral spines immediately postero-lateral to the external teeth in some specimens, but these spines are often absent from one of both sides, particularly in large specimens, possibly due to damage. Tergites: emargination of posterior borders of large tergites variable, particularly that of T.14 which may be quite markedly emarginate or may be almost straight; posterior angles of T.5 rounded or blunt, those of T.8 blunt or angulated sometimes with traces of projections, those of T.10 blunt or angulated often with slight projections, those of T.12 sometimes blunt but usually angulated with slight projections, those of T.14 blunt or angulated, sometimes rounded; posterior angles of T.4 usually rounded, sometimes very slightly projecting. Posterior projections on T.6 always well-developed, usually blunt, sometimes sharp; those on T.7 well-developed, sometimes very broad and short, with or without some sinuosity of their internal borders; those on T.9, 11 and 13 well-deVeloped with internal borders more or less sinuous; in some specimens this sinuosity is so marked, particularly on T.11 and 13, that the tip of the projection is narrow and pointed as in the figured immature female of
E. litoralis
(Fig. 12). Posterior border of intermediate tergite sinuous in male, with rounded or trapezoidal emargination in female. Coxal pores: 35 to 65 in four to six irregular rows on each of the 12th to 15th coxae, most numerous on 14th. Glandular pores of 15th legs (Fig. 4): concentrated on all aspects of femur, tibia, tarsus and metatarsus, and also on internal and dorsal
aspects
of prefemur as figured by Verhoeff (1937: 172, fig. 2) for "
Polybothrus fasciatus
".
Chaetotaxy of 14th and 15th legs
General setae: on the metatarsus sparse and short, not more than a quarter the diameter of the article in length, usually much less in males (Fig. 5), sometimes a little longer in females; on the tarsus, tibia, femur (Fig. 4) and prefemur shorter and sparser than on metatarsus, sometimes a little longer in females than in males. Seriate setae (Figs. 5 & 6): as in
E. fasciatus
, but in females those of the 15th metatarsus may be reduced to about three at the distal extremity of the article. Spinous setae: on the 14th tarsus the ventral external and ventral internal setae are of much the same structure (Fig. 6) and correspond to the
"spines"
VaTa and VpTa described by Brolemann (1930: 246) in "
Bothropolys fasciatus
"; in addition to the ventral internal seta there are, in large specimens, a few more spinous setae at intervals along the ventro-internal border of the tarsus. On the 13th and more anterior legs the spinous tarsal setae are similar to those on the 14th but rather more numerous along the ventro-internal border; the tibia of each of these legs also bears a linear series ofspinous ventro-internal setae, the most distal of which corresponds to the
"spine"
VpT of Brolemann. Same of tuft (males): short and few, but present in all adults examined (Fig. 4)
Sculpturg of male 15th legs (Fig. 4)
Prefemur with dorsal sulci distinct, internal one continuous with basal femoral pit, external one not reaching the distal end of article; basal femoral pit shallow, occupying about a quarter to a third the diameter of the base of femur, continuous distally with the narrow internal femoral sulcus which runs to the margin of the pore-free area and is of the same width and depth throughout; external femoral sulcus similar in width, starting level with the basal pit and extending further distally than the internal sulcus; pore-free area occupying distal one third or less of the internal aspect of femur, with no swelling, almost glabrous, the minute setae on its surface being just as sparse as on the rest of shaft.
Sculpturing of male 14th legs
Internal and external dorsal sulci usually fairly distinct on both prefemur and femur.
The 14th leg of
holotype
The femur (R. only) is unusually broad so that the 14th leg appears stouter than the 15th; this appearance is not due to defective development of the 15th leg as suggested by Latzel (1880: 48).
Spinulation
| Ventral |
Dorsal |
| C |
t |
P |
F |
T |
C |
t |
P |
F |
T |
| 14 |
- |
m |
amp |
amp |
am |
a |
- |
amp |
a-p |
a-p |
| 15 |
- |
m |
amp |
am |
am |
a |
- |
amp |
p |
p |
14 VpF and 15 DpT may be absent; 15 VpF and 15 DaF may be present; no coxolateral spines; a well-developed 15th accessory apical claw.
Genitalia
.
Male: posterior border of genital sternite with a median notch, more distinct in large specimens, and long marginal setae on either side, the medial setae adjacent to the notch being often shorter than the more lateral setae but showing no sharp differentiation; gonopods long and slender, basal article less than half the length of distal article. Female: two cylindro-conical spurs on each gonopod separated from one another at their insertion by about their own diameter, the internal pair being often rather smaller than the external pair even in mature specimens; claw of gonopod sharp, without denticles; dorso-lateral setae of gonopod short and stout, in an irregular band of about six setae on the first article, about twelve on the second and none on the terminal article. In large specimens the spurs of the gonopods may be short and blunt and the claw blunt, possibily due to wear and tear.
IMMATURE STADIA. There is sufficient available material to describe the last larval stadium and five further stadia which probably cover the complete post-larval lifehistory. It is difficult to make an exact comparison between each of these stadia and those described by Verhoeff (1905) for
Lithobius forficatus
;
Verhoeff's
terms are only roughly applicable and are therefore placed in parenthesis.
Fourth larval stadium
Length: about
9 mm
. Antennae: 19 to 24 articles. Prosternal teeth: usually 6+6, sometimes 6+5 or 6+4. Tergites: posterior border of last tergite (T.12) deeply emarginate; posterior projection on T.6, 7, 9 and 11 well-developed. Coxal pores: one on 12th coxa.
First post-larval stadium (agenitalis 1)
Length: about
10 mm
. Antennae: broken. Ocelli: 1+2, 1. Prosternal teeth: 6+6 or 6+7. Tergites (Fig. 7): posterior borders of large tergites deeply emarginate; posterior projections on short tergites well-developed and much narrower than in adults. Coxal pores: 2, 2, 2, 1. 14th and 15th legs: missing.
Second post-larval stadium (agenitalis 2)
Length: about
12 mm
. Antennae: broken. Ocelli: 1+2, 2. Prosternal teeth: 6+6 or 6+7. Tergites 2 as in last stadium. Coxal pores: one relatively large external pore and 4 to 6 much smaller pores on each of the 14th to 15th coxae. 14th and 15th legs: missing. Genitalia: undeveloped.
Third post-larval stadium (immaturus)
Length:
12 to 14 mm
. Antennae: 38 articles. Ocelli: 1+3, 3, 2. Prosternal teeth: 6+6 to 7+7. Tergites: posterior borders of large tergites as in last stadium; posterior projections on short tergites rather less narrow; posterior border of intermediate tergite straight. Coxal pores: 7 to 10 on each of the 12th to 15th coxae. General setae of 14th leg: those of metatarsus exceeding the diameter of the article in length. Seriate setae of 14th leg: as in adult. Spinons setae of 14th leg: not fully
developed
. Spinulation of 14th legs: as in adult. 15th legs: missing. Male genitalia: posterior border of genital sternite without a notch, with or without one or two marginal setae on either side; gonopods appears as unsegmented slender buds. Female genitalia: gonopods small with indefinite segmentation and neither spurs nor claw.
Fourth post-larval stadium (praematurus)
Length:
15 to 18 mm
. Antennae: 39 or 40 articles. Ocelli: 1+3, 3, 3 or 1+3, 3, 2, 2. Prosternal teeth 6+6 to 7+7. Tergites: approaching the shape found in adults, but posterior borders of large tergites tend to be more deeply emarginate and posterior projections on short tergites tend to be narrower and sharper; posterior border of intermediate tergite straight or slightly emarginate in either sex. Coxal pores: 10 to 19 on each of the 12th to 15th coxae, usually many more on 12th than on 15th. Glandular pores of 15th legs: as in adult. General setae of 14th and 15th legs: much longer than in adult, those of metatarsus being about as long as the diameter of the article. Seriate setae of 14th and 15th legs: those of 14th tarsus and metatarsus as in adult; those of 15th metatarsus reduced to a few at the distal extremity of the article. Spinous setae of 14th leg: not fully developed. Setae of tuft (male): absent. Sculpturing of 15th legs: indistinct in males as in females. Spinulation of 14th and 15th legs: as in adult. Male genitalia: posterior border of genital sternite with or without a trace of median notch, with about 3 or 4 marginal setae on either side; basal article of gonopod as long as distal article. Female genitalia: gonopods fairly well-developed, completely segmented, with or without minute spurs, with a small claw.
Fifth post-larval stadium (pseudomaturus)
Length:
19 to 25 mm
. Antennae: 40 to 49 articles. Ocelli: 1+3, 4, 3 to 1+4, 4, 3, 2. Prosternal teeth: 7+7 to 8+9. Tergites: as in last stadium but posterior border of intermediate tergite very slightly sinuous in male, with rounded or trapezoidal emargination in female. Coxal pores: 20 to 35 on each of the 12th to 15th coxae. General setae of 14th and 15th legs: those of metatarsus up to half and those of tarsus up to one third of the diameter of the article in length; others relatively shorter than in last stadium but longer than in adult. Seriate setae of 14th and 15th legs: as in adult but sometimes slightly reduced on 15th metatarsus. Spinous setae of 14th leg: as in adult, or the ventral external seta more slender. Setae of tuft (male): absent. Sculpturing of 15th legs: femoral sulci more distinct in males than in females, but basal femoral pit and pore-free area absent or ill-defined. Male genitalia: posterior border of genital sternite usually with a feeble median notch and about 6 to 8 marginal setae on either side; basal article of gonopod about half the length of distal article. Female genitalia: gonopods with small unequal spurs and a well-developed claw.
DISCUSSION. This is the species which, ever since the publication of
Pocock's
(1890) Synonymy (see p. 294), has been regarded by most authors as the typical form of
Lithiobius
fasciatus
Newport.
But Chamberlain (1925) designated
L.
grossipes
C. L.
Koch
as the
type
species of
Eupolybothrus
and only he, among modern authors, uses
C
. L.
Koch's
name either because he was not familiar with the current European literature or because he regarded
Newport's
description of
L. fasciatus
as inadequate.
It seems that when L. Koch redescribed
L. grossipes
in 1862 he had only the
holotype
before him and identified his other available specimens of the species either as
L. montanus
C. L. Koch
or as a
new species
,
L. festivus
, based on immature examples. Most of the material in the Koch Collection was probably named subsequent to 1862 when L. Koch must have realised the identity of these three forms with one another and labelled them "
L. grossipes
" of "
L. grossipes
,
festivus
." His reason for the continued use of the name
"
festivus
"
on some of the labels is quite obscure and was not based on immaturity.
E. grossipes
has been fairly adequately described under either
grossipes
or
"
fasciatus
"
by a number of authors. Verhoeff (1937: 178) describes T.8, 10, 12 and 14 as having rounded posterior angles and T.9, 11 and 13 as having posterior projections with almost rectilinear borders. Whereas a few of
Koch's
specimens have rounded angles on the large tergites as figured by Verhoeff (1937: 174, fig. 7, oben) for "
fasciatus
", the majority of
Koch's
and all
Pocock's
specimens have sharp or even slightly produced angles, particularly on T.10 and 12, much as figured by Verhoeff (1937: 174, fig. 7, unten) for
Polybothrus baldensis
. The posterior projections on the short tergites are more often as described and figured by Verhoeff (1937: 173, fig. 6) for
"
fasciatus
"
but there is considerable variation and those of T.11 and 13 may have their internal borders so sinuous and emarginate, and their extremities so narrow, that they resemble the figure of
E. litoralis
(Fig. 12) or even
Verhoeff's
(1937: 173, fig. 4) figure of T.13 in
P. baldenis
. The exact shape of the trunk tergites is so variable that it is of little use as a taxonomic character, and the above description as well as the figures of
E. litoralis
(Figs. 10 & 12) might apply equally to each of the three species under consideration.
But this tendancy of the tergites to differ from the shape described by Verhoeff for
"
fasciatus
"
does occur more often in immature specimens both as regards the narrowing of the projections on the short tergites and the sharper angulation of the large ones; sharper angulation of the latter is often associated with increased concavity or emargination of their posterior borders as described by Verhoeff (1934: 72) for
Polybothrus fasciatus albanicus
.
Verhoeff's
statement, therefore, when describing
albanicus
, that tergal projections are weaker (blunter) in immature than in mature specimens is not true of
E. grossipes
although it may be true of many species of
Lithobiidae
. These variations are well shown by
Pocock's
specimens, the majority of which are immature. For example, angulation and emargination of T.14 is most marked in the earliest post-larval stadium (Fig. 7) and the last tergite (T.12) of the fourth larval stadium is even more deeply emarginate.
Although in mature males the seriate setae of the 15th metatarsus are extensive (Fig. 5) and the general setae of this and other articles of the 14th and 15th legs are very short, in immature specimens (and sometimes in mature females) the seriate setae tend to be reduced in number and the general setae are relatively longer. There actually seems to be an inverse relationship between the length of the general setae relative to the diameter of the article on which they are borne and the degree of maturity of the specimen.
We
thus have two sets of characters, the shape of the tergites and the length and arrangement of the setae on the legs, particularly those of the 15th metatarsus, which are dependant to a very great extent on degree of maturity. The validity of such species and subspecies as
P.
baldensis
Verhoeff (1937)
,
P. fasciatus
albanicus
Verhoeff (1934)
and
P. fasciatus
storkani
Verhoeff (1934)
, all depending on one or both of these characters for their definition, is therefore open to question.