The Bijdrgen tot de Dierkunde, 55 (1): 67-77 1985 Modified mouthprts in hydrophilous cve millipedes (Diplopod) by Henrik Enghoff Zoologicl Museum, University of Copenhgen, Universiletsprken 15, DK-2100 Copenhgen, Denmrk Abstrct unrelted cve-dwelling In severl millipedes from southern Europe nd the Cucsus, the mouthprts re convergently modified: The biting/msticting ofthe prts mndibles re reduced, wheres the pectinte lmelle re hypertrophied; the medil lbrl teeth re often reduced, nd the gnthochilrium often broder thn usul. Species with modified mouthprts re known in the Leucogeorgi, fmilies Julide (gener Trogloiulus, Typhloiulus), Blniulide (genus Vscoblniulus), nd Polydesmide (genus Serrdium). Julid nd blniulid The mteril studied derives from the following museums: MNV: Museo Civico di Stori Nturle, Veron, Itly. MSNB: Museo Civico di Scienze Nturli, Bergmo, Itly. NHMS: Ntionl Nturl History Museum, Sofi, Bulgri. NHMW: Nturhistorisches Museum, Vienn, Austri. ZMUC: Zoologicl Museum, University of Copenhgen, Denmrk. ZSBS: Zoologischc Smmlung des Byerischen Sttes, Munich, Germny (F.R.G.). species with modified mouthprts tend to hve fewer segments thn relted species. Avilble evidence suggests tht species with modified mouthprts live in, or t the edge of, subterrnen wter with bodies. It is tht the modified mndibles suggested enlrged pectinte lmelle function s kind of their JULIDAE Leucogeorgi filter, screening suspended orgnic mteril from the Mteril exmined. type-series of L. longipes wter. The shortened body my be consequence of lessened need for pushing power, nd perhps endows the species with higher speed of locomotion. Verhoeff, 1930 (ZSBS). Verhoeff (1930) described the new genus nd species Leucogeorgi longipes from cve in the Cucsus. The species devited from ll other INTRODUCTION Cves hve been invded independently by numerous unrelted lineges of millipedes. julids known t tht time in severl mouthprt chrcters: the lbrum lcks the usul three medin teeth; the biting/msticting of the prts mndibles, viz., the externl nd internl teeth, Mny of the cve-dwelling species differ from nd the molr plte, re reduced, wheres the their cpigen reltives in being blind, pler, longer, nd hving longer legs nd ntenne, mndibulr pectinte lmelle re hypertrophied (terminology from Enghoff, 1979); the but otherwise they do not show morphologicl significnt devitions. It hs been known for long time tht few of the cve-dwelling Julide re distinguished by set of prticulr gnthochilrium is broder thn usul, the lterl plps re enlrged, the lingul lobes very brod, nd the sete of the lmelle lingules re A mostly picl. further modifiction, not modifictions of the mouthprts. Prompted by noted by Verhoeff, ffects the posterior the discovery of such species belonging to the hypophryngel node, so chrcteristic of the genus Trogloiulus, I reviewed the literture per- Julide (Enghoff, 1981), which is reduced in tining to this phenomenon, with some interesting results, which re reported on here. Leucogeorgi (like in Typhloiulus usugi Mnfredi, 1953, T. edentulus Attems, 1959, nd Trogloiulus Similr cses of mouthprt modifiction in the binii, fig. 4). fmilies Blniulide nd Polydesmide re lso Verhoeff ws so the modified impressed by discussed. mouthprts (s such mouthprts will be
A The lmost with One Numerous 68 PROCEEDINGS 6th INTERNATIONAL CONGRESS OF MYRIAPODOLOGY termed in the present pper) tht he (1926-32) Modified mouthprts in the genus Typhloiulus erected seprte fmily to ccommodte Leucogeorgi. The Leucogeorgiide did no survive for long: Lohmnder (1936) demonstrted tht Leucogeorgi hs reltives, viz., species of the genus Archileucogeorgi Lohmnder, 1936, were described by Strsser (1962, b), s occurring in T. (Speleoblniulus) serbni (Ceuc, T. edentulus 1956), (Attemsotyphlus) Attems, 1959, nd T. (Stygiiulus) usugi Mnfredi, 1953. Strsser did not mention detils of the mn- with perfectly norml julid mouthprts. The dibles in T. usugi, but I found them, like those two gener re identicl in gonopod mor- of T. serbni nd T. edentulus, to be of the phology nd re both currently clssified in the Leucogeorgi type one notble ddition, Lohmnder described Julide. (1936) two species of Archileucogeorgi from the Cucsus, of which one ws found in cve. Golovtch (1981, 1983, nd in litt.) reports on two further Leucogeorgi both nd severl spp., troglobites, nmely tht there re five insted of the usul four pectinte lmelle. Typhloiulus (Inversotyphlus) longipes Strsser, 1973, my represent n intermedite stge in the evolution of Leucogeorgi- like mouthprts: new Archileucogeorgi spp., mostly troglobites, The clypeus nd gnthochilrium re brod from the Cucsus. (Strsser, 1973), but the mndibles nd hypophrynx re unmodified (pers. obs.). Trogloiulus nd Typhloiulus Mteril exmined. type-series of Trogloiulus binii Enghoff, 1985 (MSNB), nd numerous specimens of the "norml" Tr. boldorii Mnfredi, 1940, Tr. mirus Mnfredi, 1931, nd Tr. minimus Mnfredi, 1935 (MSNB nd BLANIULIDAE Vscoblniulus cbidochei MNV). mle topotype of Typhloiulus usugi Mnfredi, 1953 (MNV). The holotype of T. edentulus (NHMW). A Mteril exmined. O", one 9, topotypes femle nd juvenile prtype of T. longipes Strsser, 1973 (ZMUC, Muries ded.). "norml" (NHMS). Specimens of the "norml" T. montellensis blniulids. Verhoeff, 1930, T. psilonotus (Ltzel, 1884), T. strictus (Ltzel, 1882), nd T. illyricus stygis Verhoeff, 1933 (ZMUC, MSNB, nd ZSBS). Wheres the modified mouthprts re currently Muries described the (1967) new genus nd species Vscoblniulus cbidochei from the fmous cve "Gouffre de l Pierre-Sint- Pyrenen Mrtin". V. cbidochei differs from the genus-defining in the cse of Leucogeorgi nd numerous other cvernicolous blniulids Archileucogeorgi, this is not so in the Itlin- Blkn gener Trogloiulus nd Typhloiulus, (gener Blniulus, Occitniulus, Euzkdiulus, etc.) in severl mouthprt chrcters: The lb rum is which both belong in the somewht dubious brod nd bulging nd hs the medin teeth tribe Typhloiulini (Strsser, 1962). strongly reduced; the mndibles hve bout Strsser (1977) reviewed the North Itlin dozen pectinte lmelle insted of the usul genus Trogloiulus Mnfredi, 1931, t tht time consisting of six species exclusively troglobites, nd ll with norml mouthprts. But in 1978, Strsser described seventh species from cve in Bresci: Tr. viltii, with four; the gnthochilrium is comprtively short, nd hs the externl distlly enlrged, plps very strong (Muries, 1967). I hve exmined mndible of mle V. cbidochei with scnning electron microscopy mouthprts entirely of the Leucogeorgi type. A nd found some further modifictions (figs. second Trogloiulus with modified mouthprts (Tr. binii) ws described by Enghoff (1985). 8-10). The externl tooth seems to be bsent ltogether; the internl tooth nd the molr Figs. 1-7 show scnning electron microgrphs plte re reduced; the teeth of the pectinte of importnt detils in the mouthprts of this lmelle (11 lmelle in the mndible exm- species, compred boldorii Mnfredi, 1940. with the "norml" Tr. ined) re extremely long, slender, nd densely set.
- 1985 BIJDRAGEN TOT DE DIERKUNDE, 55 (1) 69 Figs. 1-5. Mouthprts: 1, 3, 4, Trogloiulus binii; 2, 5, Tr. boldorii, norml, troglobiont julid. (1-2, left mndible, gnthl lobe; 3, externl nd internl teeth of right mndible; 4-5, posterior node of hypophrynx). Scles 100 μm (1, 2, 5), 50 μm (3, 4).
70 6th PROCEEDINGS 6-7. Figs. Pectinte Most of prllels lmelle: these The tinte lmelle, deserves the s Trogloiulus binii; modifictions those found in the to bove. view 6, strongly possession from is four unique 1981) (rrely five) CONGRESS boldorii. exct discussed pec- nd systemtic point Enghoff (1979, of re julids Tr. 7, enlrged number of however, comment well: INTERNATIONAL of found tht pectinte Scles 20 lmelle OF MYRIAPODOLOGY μm. insted synpomorphy Blniuloide V. + of for hypothesis number higher julid (nd to it be is + potent bove-cited quite ppers). Juloide. flsifier of embrrssing tht I overlooked Muries' when I report the is superfmilies Nemsomtoide cbidochei ppers this the Severl wrote other
these BIJDRAGEN TOT DE DIERKUNDE, 55 (1) - 1985 71 Vscoblniulus Figs. 8-11. Mndibulr gnthl lobes with pectinte lmelle: 8-10, cbidochei; 11 norml blniulid. Scles 100 μm (8), 50 μm (9), 20 μm (10, 11). Nopoiulus kochii (Gervis), chrcters, however, indicte tht the lrge blniulids s well probbly re cses of number of lmelle is secondry modifiction: V. cbidochei exhibits severl of the blniulid secondry reduction. The of ocelli rrngement escpes evlution in this blind species. pomorphies mentioned by Enghoff (1981): nterior mle legs with leflike sete, penis The enlrged number of pectinte lmelle is single, nterior gonopods without flgell, prllelled, lthough on much smller scle, vulvl invgintions deep. The chrcteristic blniulid mndibulr forceps is bsent, but so it is in severl other txonomiclly scttered in Typhloiulus usugi where the number of lmelle (five) is still within the "llowble" limits.
PROCEEDINGS 6th INTERNATIONAL CONGRESS OF One 72 MYRIAPODOLOGY POLYDESMIDAE Leucogeorgi hve 49-54 segments (Golovtch, 1983 nd in litt.). Archileucogeorgi spp. hve 36 - Serrdium sp. c. 60 segments (Lohmnder, 1936, Golovtch, Mteril exmined. juv. 9 (17 segments) of in litt.). Serrdium from "Grott sp. dell'acqu" del Ponte de Vej, S. Ann d'alfedo, Itly, 16.IV. 1983, leg. Coduro (MNV). Specimens of the "norml" polydesmids S. Vscoblniulus cbidochei (fig. 18) hs 40-44 nd thus lies in the lower end of segments the rnge covered other cvernicolousblniulids by V. (Brolemnn, 1923; Demnge, 1981). cbidochei is further chrcterized by prticulrly stout legs. The shortbodied, nd Serrdium sp. Polydesmide re generlly from "Grott dell'acqu" bers no signs of further shortening (only one juvenile known, however). hirsutipes nd S. longicorne (ZMUC, Pisoni nd Osell ded.). In mteril of cvernicolous millipedes from northern Itly I found single juvenile specimen of the genus Serrdium, in which the pectinte lmelle re remrkbly enlrged, nd the externl nd internl teeth diminished, in comprison with the S. hir- "norml"species sutipes Verhoeff, 1941, nd S. longicorne (Silvestri, 1894) (figs. 12-15). Lbrum nd gnthochilrium of the specimen re identicl to those of S. hirsutipes nd S. longicorne. This is, to my knowledge, the first recorded instnce of "leucogeorgioid" mouthprt modifiction in the order Polydesmid. HABITAT The bionomiclly best known of the species with modified mouthprts is Trogloiulus viltii. Strsser cites D. (1978) Vilti, who wrote (trnslted from Itlin): "The millipedes were BODY SHAPE The typhloiulines with modified mouthprts re collected in n ctive mender in the bovementioned cve, clled 'brnch of the wterflls' becuse of the presence of mny wterflls. The wterflls chnge, in periods of smll remrkbly short-bodied compred with con- flow, into film of wter running over the ver- generic species. Adults of Typhloiulus edentulus hve 33-38 segments (Attems, 1959; Strsser, ticl prts of the cnl excvted in the rock... When collected, the dult diplopods were 1962b; the record of 58 segments by Attems heding upstrem, covered by t lest couple (1951) is erroneous), T. usugi lso hs 33-38 of millimeters of wter, wheres the juvenile segments (Mnfredi, 1953; Strsser, 1971), nd stdi could be found t the sides, where only T. serbni hs 29-35 segments (Ceuc, 1956, the spry ws reching. The ir temperture 1961). T. longipes, with only prtly modified mouthprts, hs 37-41 segments (Strsser, ws 8.7 C, the wter temperture 7.5 C". Limited informtion is vilble on the other 1973). Other species of Typhloiulus hve 39-79 julids with modified mouthprts, but wht there segments (Strsser, 1962). ( T. stregi Strsser, hs norml 1973, mouthprts nd 32-40 segments. It is, however, not typhloiuline is suggests tht they my live like T. viltii: Leucogeorgi longipes ws collected in puddles (Verhoeff, 1930). L. rediviv Golovtch, 1983, (pers. obs.).) nd L. n. sp. inhbit "very humid prts of the viltii Trogloiulus hs 30-34 segments Tr. binii (Strsser, 1978), (fig. 17) hs 27-30. cves" (Golovtch, in litt.). T. serbni ws found "prtly on the cve wll, prtly on stones Other Trogloiulus spp. hve 41-87 segments ner subterrnen wter. Both wlls nd stones (Strsser, 1977) (fig. 19). The difference is less cler-cut in Leucogeorgi nd Archileucogeorgi: L. longipes with 28-34 were wet" (Ceuc, 1956, trnslted from Rumnin). No hbitt informtion is vilble on T. edentulus nd the nominte segments (fig. 16) hs the sme body shpe s forms of T. usugi, but T.. gentine ws found the modified typhloiulines, but other species of "on wet, clyey... wlls of the cve" (Strsser,
- 1985 BIJDRAGEN TOT DE DIERKUNDE, 55 (1) 73 Figs. 12-15. Left mndibulr gnthl lobes: 12-13, Serrdium sp. from Grott dell Acqu ; S. 14-15, longicorne. Scles 50 μm (12), 100 μm (13-15).
... nd = 74 PROCEEDINGS 6th INTERNATIONAL CONGRESS OF MYRIAPODOLOGY Figs. 16-19. Body shpe of species with modified mouthprts (16-18), compred with norml troglobiont julid (19): 16, Leucogeorgi longipes; 17, Trogloiulusbinii; 18, Vscoblniulus cbidochei; 19, Trogloiulus boldorii. Robert Nielsen del. Scle 5 mm. 1971, trnslted from Germn), nd T.. fimbrtius ws found in cve bout which Strsser (1971) wrote (trnslted from Germn): "The remining fun is never found together with Aphenops [ genus of crbid beetles] nd other encountered species which spring cve, which is known nd often come to bits. (This is in contrst to wht one visited becuse of its built-in old church, hs cn usully see in numerous cves t low been often nd thoroughly investigted, without Typhloiulus hving been found before. Its rrity might be the explined by fct tht the cve is sometimes flooded". The cve in which Trogloiulus binii ws collected is described by ltitude in the sme region, where diplopods of the genus Typhloblniulus [ Blniulus] litterlly Bini (1977) who further informs (R. Pisoni, in swrm on bits.) This new genus described by Muries is thus quite distinct: in ddition to morphologicl criteri its ecology is quite different becuse it is the only species litt.) tht the smpling sites for the species re lwys very muddy, becuse they re often flooded. Vscoblniulus cbidochei fits into this picture too (Muries, 1967; Cbidoche, 1968): which frequents the bnks of subterrnen wtercourses..." (Cbidoche, 1968: 670, trnslted from French). The Serrdium from "Grott dell'acqu" "Among the species living in this cvity, nicely rounds off the pttern of hydrophily: it ws collected under stones in the bed of the little Vscoblniulus cbidochei should be especilly emphsized: it is loclized very differently from the brook of the cve (G. Osell, in litt.).
Typhloblniulus) BIJDRAGEN TOT DE DIERKUNDE, 55 (1) - 1985 75 HABITAT OF RELATED SPECIES DISCUSSION Only scnt hbitt informtion on "norml" The foregoing sections indicte n interesting could be extrcted from cve julids the literture. This in itself my indicte tht these species do not occur in very specil hbitts, unlike t lest some of the with species modified mouthprts. Golovtch (in litt.) reports on "mny new "syndrome". The ffected species hve their mouthprts modified; in prticulr, the biting prts of the mndibles re reduced, wheres the pectinte lmelle re hypertrophied. The species tend to be short-bodied, nd vilble hbitt informtion suggests tht most, ifnot ll Archileucogeorgi inhbiting very wet cves... of them, fvour very wet hbitts nd might However, some Archileucogeorgi hve been found on reltively dry wlls in cves, nd some even in the forest litter (epigeiclly)". deserve the designtion semiqutic. Relted species with norml mouthprts (nd longer bodies) pper occupy other, less wet, Trogloiulus minimus hs been collected "on hbitts in the cves ( Typhloiulus illyricus stygis rotting wood" nd "under wter collector seems to be n exception. I hve exmined this (sewer)" (unpublished mteril in MSNB, Enghoff det.). Typhloiulus bureschi Verhoeff, 1926, hs been found on moist nd in bt cly guno (Gueorguiev & Beron, 1962). T. strictus (Ltzel, 1882) occurs in ccumultions of decying plnt nd niml wste (Negre & T. Negre, 1979). species nd scertined tht its mouthprts re norml.) It is remrkble tht the syndrome recurs in three fmilies nd tht within the Julide, it hs pprently evolved severl times independently: The Leucogeorgi, Typhloiulus, nd Trogloiulus species with modified ll hve close mouthprts lbnicus Attems, 1929, ws found under stones reltives with norml mouthprts. The three t cve entrnce (Strsser, 1962). T. illyricus Typhloiulus species involved re even clssified Verhoeff, 1929, ws found bundntly on n in three different subgener! ccidentlly introduced shef in Slovenin cve (Strsser, 1966), but T. illyricus stygis ws collected on moist stlgmites nd on cly nd POSSIBLE FUNCTIONAL SIGNIFICANCE wood in very wet cve with cly nd mny muddy puddles (Verhoeff, 1933). Brolemnn writes bout the (1923: 411) hbitt of cvernicolous blniulids tht species of Blniulus ( = nd Archichoneiulus hve been collected in bundnce on bt guno, on stlgmitic flows nd on Modified mouthprts The function of julid mouthprts ws described by Fechter nd (1961), Mnton (1964) described the mndibulrfunction of species belonging to the relted order According to these uthors, the externl nd Spirostreptid. internl decomposing vegetble nd niml debris. teeth of the mndibles pick up food prticles They hve lso been found eting old drops of which re then ground between the molr sterin (from the cndles of speleologists) mould growing upon them. (See under Vscoblniulus cbidochei.) or lso bove Serrdium hirsutipes nd S. hve been longicorne collected in numerous North Itlin cves, some with, others without wter. (G. Osell, R. In the pltes. of cse the Julide, molr pltes grind the food ginst the node hypophryngel (which is chrcteristic of the Julide nd one further fmily, the Trichoblniulide, see Enghoff, 1981). According to Fechter (1961), the pectinte lmelle lso ssist in the picking Pisoni, in litt.). The only vilble exct hbitt up nd grinding of food prticles. (Verhoeff notes, however, concern n undescribed Ser- (1926-32: 138) interpreted the pectinte rdium species with norml mouthprts, which ws found in gret numbers on the wlls nd on vrious orgnic mterils (wood, etc.) in n rtificil cve ner Bergmo (R. Pisoni, in litt.). lmelle s grooming orgns; however, B. D. Vlentine (in litt.), who hs studied grooming in 14 fmilies of hs never seen the millipedes, pectinte lmelle being used for grooming.)
76 PROCEEDINGS 6th INTERNATIONAL CONGRESS OF MYRIAPODOLOGY species of the order Polydesmid. A correltion The medin lbrl teeth re used s n nchor during burrowing (Fechter, 1961). The modifictions described in the preceding between hydrophily nd high speed of locomotion is thus suggested. sections must obviously ffect these functions profoundly. Verhoeff (1930) hypothesized tht the enlrged pectinte lmelle of Leucogeorgi scrpe off prticles of some soft substnce present in lrge quntity. He suggested this substnce to be either fine, moist ooze or more or less decyed guno. Strsser (1978) found tht severl specimens of Trogloiulus viltii hd their preorl cvity stuffed with some extrneous substnce in such tht quntities it prevented them from their "closing mouth". He suggested tht the specimens might hve been cptured while "grzing" under wter nd tht the substnce might be but hrdly bt guno. ooze, I found the gut contents of Trogloiulus binii to consist of extremely finegrined mteril which could not be identified, not even under the electron scnning microscope. Bsed on this scnt evidence I cn only follow Verhoeff in hypothesizing tht these species feed on some soft, fine-grined mteril. The indictions for hydrophily led to the suggestion CONCLUSION The evidence presented here, lthough frgmentry nd insufficient, clerly indictes tht the "modified mouthprts syndrome" constitutes significnt feture in the dpttion of cve-dwelling millipedes. Tht there is prticulr functionl spect of the modifictions (probbly dpttions to the recovery of suspended food in subterrnen wterbodies) is indicted by the txonomic isoltion of the species ffected. So fr, the phenomenon hs been found only in quite restricted re: from northern Itly over the Blkn countries to the Cucsus. The reson for this be either tht my cve millipedes of other regions re less well-known, or tht they belong to tx whose mouthprts re not pre-dpted for this kind of modifiction. ACKNOWLEDGEMENTS tht the enlrged pectinte lmelle function s kind of filter. The reduction of the lbrl teeth be my correlted with non-burrowing hbits of the species involved. The gnthochilril modifictions escpe evlution t present but certinly hve functionl spect, since they re prlelled in severl unrelted lineges. Specimens for the present study were loned nd/or donted, nd useful unpublished informtion ws provided by: P. Beron H. Fechter S. (Sofi), (Munich), Golovtch (Moscow), J. Gruber (Vienn), J.-P. Muries (Pris), G. Osell (Veron), R. Pisoni (Bergmo), nd B. D. Vlentine (Columbus, Ohio). R. Nielsen nd B. W. Rsmussen took in the elbortion prt of illustrtions, nd M. Bevort nd I. Grosz trnslted Itlin nd Rumnin texts, respectively. I extend sincere thnks my to ll these persons. Body length A lrge number of body segments increses the pushing power millipede is ble to exert (Mnton, 1954). The possibly hydrophilous julids nd blniulids with modified mouthprts probbly hve little need for The burrowing. comprtively short body with long legs, quite reminiscent of certin Prdoxosomtide (order Polydesmid) probbly results in higher speed of locomotion, when with the compred relted The longer species. very strong legs of Vscoblniulus cbidochei lso recll certin REFERENCES ATTEMS, C., 1951. Neue Hohlen-Myriopoden, gesmmelt Professor Absolon. von Anz. ost. Akd. Wiss., (mth.-nturw. Kl.) 88: 253-257., 1959. Die Myriopoden der Hohlen der Blknhlbinsel. Annln. nturh. Mus. Wien, 63: 281-406, Tf. 8-9. BINI, A., 1977. Ntur in Lombrdi. Le Grotte (Ed. Regione Lombrdi, Milno). BROLEMANN, H. W., 1923. Blniulide. Myripodes (premiere serie). Archs. Zool. 61: exp. gen., 99-453, pis. I-XVI.
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