Eupolybothrus (E.) grossipes (C. L. Koch)

E. H. Eason, 1970, 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), Bulletin of the British Museum (Natural History), Zoology 19, pp. 289-310 : 295-303

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 [ Isère, 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

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.