47
THE MILLIPEDE TRIBE APFELBECKIINI
A revision of the millipede tribe Apfelbeckiini Verhoeff, 1900
(Diplopoda: Callipodida: Schizopetalidae)
PAVEL STOEV & HENRIK ENGHOFF
Steenstrupia
Stoev, P. & H. Enghoff. A revision of the millipede tribe Apfelbeckiini Verhoeff, 1900 (Diplopoda:
Callipodida: Schizopetalidae). – Steenstrupia 39 (1): 47–66. Copenhagen, Denmark. April 2008. ISSN
0375-2909.
The millipede tribe Apfelbeckiini (Callipodida: Schizopetalidae) is revised. After re-examination of
most of the type specimens of hitherto described taxa the number of valid genera and species in the tribe
is reduced to two and four, respectively. The following new synonymies and combinations are proposed
for the first time: Genus Apfelbeckia Verhoeff, 1900 = Genus Antropetalum Attems, 1926 syn.n.,
Antropetalum brazzanum Attems, 1927 = Apfelbeckia brazzana (Attems, 1927) comb.n.; Apfelbeckia
insculpta (L. Koch, 1867) comb.n. = Lysiopetalum lendenfeldii Verhoeff, 1896 syn.n.; = Apfelbeckia
enderleinii Verhoeff, 1901 syn.n.; = A. albosignata Verhoeff, 1901 syn.n.; = A. silvivaga Verhoeff,
1901 syn.n.; = A. wohlberedti Verhoeff, 1909 syn.n.; = A. hessei Verhoeff, 1929 syn.n.; = A. lendenfeldii
var. flavipes Attems, 1929 syn.n.; = A. albanica Verhoeff, 1941 syn.n.; = A. subterranea Verhoeff, 1943
syn.n.; = A. hessei var. boldorii Manfredi, 1945 syn.n.; = A. duplocalca Attems, 1951 syn.n.; = A.
lendenfeldi miraculosa Attems, 1951 syn.n.; = Karlabsolonia mirabilis Attems, 1951 syn.n. A new
species, Apfelbeckia synthesys, distinguished from its congeners by having only a single row of
pleurotergal setae and specific shape of male gonopods, is described from Olympos Mts., Greece. An
identification key to the genera and species in the tribe is provided, along with remarks on the
morphology and distribution of some species.
Keywords: Apfelbeckia synthesys sp. n., Olympos Mts., Greece, taxonomy, gynandromorphism
Pavel Stoev: National Museum of Natural History, Tsar Osvoboditel Blvd. 1, 1000 Sofia, Bulgaria. Email: [email protected]
Henrik Enghoff: Natural History Museum of Denmark, Universitetsparken 15, DK-2100 København Ø,
Denmark. E-mail: [email protected]
INTRODUCTION
The tribe Apfelbeckiini Verhoeff, 1900 (Callipodida: Schizopetalidae) is a quite homogeneous
group with regard to the shape of the male gonopods. The tribe is currently understood as containing the polytypic genus Apfelbeckia Verhoeff, 1896, and the monotypic genera Himatiopetalum Verhoeff, 1900 and Antropetalum
Attems, 1926 (Hoffman 1973). The genus Apfelbeckia is currently known to comprise 10 nominal species, and 3 subspecies/varieties of doubtful validity, widely distributed in the western part
of the Balkan Peninsula: Croatia, Bosnia and
Herzegovina, Serbia and Montenegro, and Albania. Himatiopetalum ictericum (L. Koch, 1867)
occurs on the Greek island of Corfu (Kerkira),
while Antropetalum brazzanum Attems, 1927
was hitherto known only from caves on the Dalmatian island of Brach (= Brazza) (Fig. 1).
Apfelbeckia was originally proposed as a subgenus of Lysiopetalum Brandt, 1840, with L. (A.)
lendenfeldii Verhoeff, 1896, as type species
(Verhoeff 1896). The type locality of L. lendenfeldii is the cave Bilek (Bileća) in Bosnia and
Herzegovina, subsequently it was also recorded
from other places in Croatia, Bosnia and Herzegovina, and Montenegro. Four years later, Apfelbeckia was elevated to full generic rank (Verhoeff 1900a). Later on, Verhoeff (1901) described three further species in the genus, A.
enderleinii (Bosnia and Herzegovina), A. albosignata (Dalmatia, Herzegovina and Montenegro) and A. silvivaga (Herzegovina), and re-
Steenstrupia 29(1): 47–66.
Stoev & Enghoff rettet.pmd
47
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P. STOEV & H. ENGHOFF
48
Fig. 1. Distribution of the species of Apfelbeckiini. Dotted line = approximate range of A. insculpta. 1. A. brazzana. 2. A.
synthesys. 3. H. ictericum.
described A. lendenfeldii. During the next 40
years he added A. wohlberedti (North Albania),
two varieties of lendenfeldii (abbreviatum and
herzegowinense, both from Herzegovina) (Verhoeff 1909a), as well as A. hessei (Croatia) (Ver-
Stoev & Enghoff rettet.pmd
48
hoeff 1929), A. albanica (Central Albania) (Verhoeff 1941), A. subterranea (Serbia), A. lendenfeldii caligulifer and A. lendenfeldii hebes (both
from Herzegovina) (Verhoeff 1943). In addition
to Verhoeff’s work on the genus, Attems (1929,
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49
THE MILLIPEDE TRIBE APFELBECKIINI
1951, 1959) described the following apfelbeckiines: A. lendenfeldii var. flavipes (North Albania),
A. duplocalca (Bosnia), A. lendenfeldi miraculosa (Bosnia and Herzegovina, Montenegro) and
Karlabsolonia mirabilis (Dalmatia). Manfredi
(1945) described A. hessei var. boldorii from
North Albania. Strasser (1971) proposed the synonymy of A. lendenfeldii caligulifera and A.
lendenfeldii hebes under A. lendenfeldii lendenfeldii, and Mršić (1994) did the same with the
subspecies abbreviatum and herzegowinense.
Strasser (1971) and Hoffman (1973) suggested
the synonymy of the genus Karlabsolonia and
the subgenus Haplobeckia Verhoeff, 1941 under
Apfelbeckia s. str.
The postembryonic development in Apfelbeckia was studied by Lang (1935), who described
eight stadia in this species. Analysing Lang’s
data, Enghoff et al. (1993) concluded that
Apfelbeckia probably develops through teloanamorphosis.
Carnivory of A. lendenfeldii was studied by
Verhoeff (1900c) (see also Hoffman & Payne
1969), as the species was reported to feed on
earthworms, flies, spiders and centipedes.
Additional faunistic and taxonomic information concerning this or that species of Apfelbeckiini can be found in the papers of Latzel
(1884, 1888), Karlinski (1894), Verhoeff (1897,
š
1899, 1909b, 1932), Kovacević
(1931), Lang
(1939), Strasser (1962, 1970), Ceuca (1964),
Tomić -Jovanović (1964), Mauriès et al. (1997),
Guéorguiev et al. (2000), Makarov et al. (2004),
and Stagl & Stoev (2005).
The overall picture gathered from the reviewed literature is that the current taxonomy of
the group is highly outdated, with all taxa lacking
adequate descriptions, often misidentified by
subsequent researchers, and types, though well
preserved and completely accessible in European
museums, having never been re-examined. Starting this project as a simple description of a new,
quite distinct species of Apfelbeckia found by a
Danish zoological expedition in Greece, during
the work the need in revising the whole tribe
became evident. Along with the description of
the new species we have re-examined the type
specimens and older collections of apfelbeckiines preserved in the Museum für Naturkunde,
Humboldt Universität zu Berlin (ZMB), Natur-
Stoev & Enghoff rettet.pmd
49
Fig. 2. Apfelbeckia synthesys sp. n., female paratype. Photo G.
Brovad.
historisches Museum Wien (NHMW), Zoologische Staatssammlung München (ZSM), and
American Museum of Natural History, New
York (AMNH). In addition to these collections,
Dr. Richard Hoffman (Virginia Museum of Natural History, Martinsville, USA) kindly placed
at our disposal his unpublished observations and
sketches of the type of Lysiopetalum insculptum
L. Koch he had made during a visit to the Natural
History Museum, London (BMNH). New collections comprising unidentified specimens were
studied from the Muséum d’Histoire Naturelle,
Genève (MHNG), Zoological Museum of the
University of Amsterdam (ZMAN), National
Museum of Natural History (Naturalis), Leiden
(NNM), the Natural History Museum of Denmark (ZMUC), and the National Museum of
Natural History, Sofia (NMNHS).
MATERIAL AND METHODS
The holotype and some paratypes of the new
species are preserved in ZMUC, male and female
paratypes are deposited in NMNHS. All specimens are preserved in 70% ethanol. The illustrations were made with a camera lucida.
28-03-2008, 13:47
P. STOEV & H. ENGHOFF
50
Abbreviations:
M – male
F – female
t. loc. – type locality
PT – pleurotergite
TAXONOMY
Family Schizopetalidae Verhoeff, 1909a
Subfamily Acanthopetalinae Hoffman &
Lohmander, 1964
Tribe Apfelbeckiini Verhoeff, 1900a, p. 50,
char. emend.
Diagnosis
Moderate-sized to large callipodidans. Females
usually larger than males. Body length ca. 50–
100 mm; diameter of midbody PTs ca. 2.5–4.8
mm. Number of PTs in adults usually 46–50.
Body colour from dark brown to brown-yellowish; usually a row of yellowish lateral spots stretching along body just below level of ozopores
(Fig. 2). PT slightly higher than broad in crosssection. Male PTs 6 and 7 not enlarged. Metazonites slightly elevated compared to prozonites;
dorsal crests either moderately (Apfelbeckia) or
well developed (Himantiopetalum); poriferous
crests normal (not enlarged like in, e.g., Caspiopetalidae or Paracortinidae). Ozopores visible
from sixth to antepenultimate PT lying either on
crest 8 or 9 (on midbody PTs). Chaetotaxy: setae
long and apically pointed, usually 8–10 on each
hemipleurite (possibly more), anterior PTs with
an anterior as well as a posterior row of setae.
Head either convex in both sexes or concave
(males of A. brazzana), usually densely covered
with minute setae. All legs very long, ending with
conspicuous claw. Coxae of all pre-gonopodal
legs without any particular modifications, tarsi
bipartite. Coxal sacs present on legs 3–16. Hypoproct either tripartite (Apfelbeckia) or undivided
(Himantiopetalum); paraprocts divided into
small dorsal and larger ventral sclerites, dorsal
sclerites with a pair of macrosetae, ventral sclerites covered with numerous macrosetae along
free margins.
Gonopods comprising modified male leg-pair
Stoev & Enghoff rettet.pmd
50
8 only; exposed ventrally beyond pleurotergal
edge; tracheal apodemes strongly expanded;
sternum either soft membrane or a chitinized,
subtrapezoidal plate lying perpendicularly between leg-pair 7 and gonopods; femoroid arising
from posterior part of coxa, divided into a large,
broad, shield-like outer branch (“Tibialabschnitt” in German literature) and a subterminal
inner branch (“Kanalast”). Inner branch ending
with solenomere and parasolenomere. Outer
branch (shield) with a very characteristic sponge-like process (“Blasiges bestacheltes Organ”)
covered with spines and lying posterolaterally on
its outer surface. In Apfelbeckia the outer branch
forms a long, anteromedial process of a characteristic boot-like shape (tarsus).
Female second leg-pair unmodified. Vulvae
extrusable, long, reaching leg-pair 8 when folded
backwards (in A. insculpta).
Remarks
Apfelbeckiini differ from the other two acanthopetaline tribes, Acanthopetalini (with three
genera, Acanthopetalum Verhoeff, 1900, Balkanopetalum Verhoeff, 1926 and Eurygyrus C.L.
Koch, 1847) and Prolysiopetalini (with a single
genus, Prolysiopetalum Verhoeff, 1909) by having male pre-gonopodal legs unmodified and by
the existence of a sponge-like organ on the outer
branch of the gonopods.
Distribution
The tribe Apfelbeckiini shows a quite compact
distribution in the western part of the Balkan
Peninsula, from the central Dalmatian coastline
in the north to Olympos Mts. in the south, and
from the Ionic island Korfu and the Dalmatian
islands of Brach, Hvar, Korchula and Vis in the
west to western Serbia and Olympos in the east.
A gap in the range existing in south Albanian
mountains and the nearest parts of Pindus Mts. in
northwestern Greece (Fig. 1) could be explained
by insufficient sampling in the region.
Genus Apfelbeckia Verhoeff, 1896
Lysiopetalum (Apfelbeckia) Verhoeff, 1896: 475. Type species: Lysiopetalum lendenfeldii Verhoeff, 1896, by monotypy.
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51
THE MILLIPEDE TRIBE APFELBECKIINI
A
C
B
D
Fig. 3. Apfelbeckia brazzana, paralectotype, gonopods. A. Anterior view. B. Posterior view. C. Anterio-mesal view. D. Lateral
view. – a, coxal process 2; g, process on inner surface of ob; h, process on t; ib, inner branch; k, sponge-like process; ob, outer
branch; p, coxal process 1; sg, seminal groove; st, sternum; t, tarsal process.
Antropetalum Attems, 1926: 180, 222. Type species: A. brazzanum Attems, 1927, by subsequent designation of Attems
(1927). Syn. n.
Haplobeckia Verhoeff, 1941: 181 (subgenus of Apfelbeckia).
Type species: Apfelbeckia silvivaga Verhoeff, 1901, by monotypy. Synonymy proposed by Hoffman (1973).
Karlabsolonia Attems, 1951: 256. Type species: K. mirabilis
Attems, 1951, by monotypy. Synonymy proposed by Strasser
(1971).
Diagnosis
A genus of Apfelbeckiini with smooth collum,
tripartite hypoproct, and outer femoroidal branch
of gonopods solid.
Stoev & Enghoff rettet.pmd
51
Distribution
Greece, Albania, Montenegro, Bosnia and Herzegovina, Croatia (including the Dalmatian islands of Brach, Hvar, Korchula, Mljet and Vis),
and Serbia.
Remarks
In the original description of Antropetalum, Attems (1926, 1927) did not clearly distinguish this
genus from Apfelbeckia. Even more, in his quite
confusing key to the genera of Schizopetalidae
he noticed that the gonopod conformation of the
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P. STOEV & H. ENGHOFF
52
new genus is similar to that of Apfelbeckia. The
characters that he used to separate Antropetalum
from Apfelbeckia, like the length of tarsi of male
postgonopodal leg-pair, shape of anal segment,
etc., are certainly of very little or no value in
Callipodida, and hardly could justify the erection
of a genus. The presence of a frontal concavity in
males of Antropetalum is the only character that
seems to genuinely separate the genus from
Apfelbeckia and Himatiopetalum. However, it
seems that in Callipodida this character is significant only at the species level, as analogues can be
found in the schizopetalid genus Prolysiopetalum Verhoeff, 1909 and the caspiopetalid genus
Bollmania Silvestri, 1896. In the latter, males
usually have a frontal knob on the head, but there
are also exceptions, e.g., Bollmania oblonga Golovatch, 1979 from Tajikistan. In B. nodifrons
Lohmander, 1933, the frontal knob exists only in
adult males, being absent in subadults (cf. Golovatch 1979, Stoev & Enghoff 2005). Examination of the type material of Antropetalum brazzanum in the NHMW confirmed that the gonopods differ only in minor details from those of
Apfelbeckia insculpta, so the erection of a genus
solely on the basis of head conformation and
other insignificant features appears unjustifiable.
Therefore, we herewith formalize the synonymy
of Antropetalum under Apfelbeckia.
Apfelbeckia brazzana (Attems, 1927) comb. n.
Figs 1, 3, 9A
Antropetalum brazzanum Attems, 1927: 114, figs 122–124.
š Brach Island,
Type locality: caves near Milna and Nerezišce,
Dalmatia, Croatia.
Antropetalum brazzanum. – Verhoeff, 1929: 634. – Attems,
192: 291, 322. – Strasser, 1971: 31. – Mršić, 1994: 238. –
Stagl & Stoev, 2005: 7, fig. 4.
Material examined:
Type specimens: 1 adult M with dissected gonopods in microtube (lectotype, by present designation); 1 further M with
dissected gonopods in microtube, 1 juv., two slides of legpairs 6–9, (paralectotypes), Inv. No. 2304 [Dalmatia, Brazza
Island, Cave II near Milna, 23.vii.1912, coll. Raab & Wettstein]; Further paralectotypes: 4 adult FF, subad. F, Inv. No.
4387 [Brazza Island, cave I near Milna, 22.vii.1912, coll.
Raab & Wettstein]; 5 adult FF, one of them mounted in a
special glass cylinder for exhibition, Inv. No. 4388 [Cave near
š (Neresi), 20.vii.1912, coll. Raab & Wettstein] (all
Nerezišce
in NHMW) (cf. Stagl & Stoev 2005).
Stoev & Enghoff rettet.pmd
52
New records: Croatia: 2 MM, 1 F, 2 juv.?, island of Hvar,
š
village of Hvar, cave Markova spilja (= Grcka
spilja),
N43º11.451 E16º24.170, alt. 68 m, 15.viii.2006, t=18ºC,
under stones, B. Petrov, S. Lazarov leg. (NMNHS); 1 M, 2
subad., same island, village of Humac, cave Grabćina spilja,
alt. 230 m, N43º08.050 E16º45.222, under stones, t=16ºC,
16.viii.2006, B. Petrov, S. Lazarov leg. (NMNHS).
Diagnosis
Well distinguished from the other species in the
tribe by the modified head in males, and from the
other congeners by the presence of an outgrowth
on the inner surface of the outer branch of the
gonopod femoroid (g in Fig. 3B).
Distribution
Croatia: Brach Island: caves near the villages of
š Hvar Island: caves near the
Milna and Nerezišce;
villages of Humac and Hvar (see also below
under A. insculpta’s distribution).
Apfelbeckia insculpta (L. Koch, 1867) comb. n.
Figs 1, 4, 5, 9B
Lysiopetalum insculptum L. Koch, 1867: 893. Type locality:
unspecified sites in Montenegro and Dalmatia.
Lysiopetalum insculptum. – Latzel 1884: 235. – Gasperini
š
1892: 8. – Attems 1929: 322. – Kovacević
1931: 71. – Stagl &
Stoev 2005: 14.
Lysiopetalum cognatum Latzel, 1884: 234. Type locality:
Ragusa (now Dubrovnik), Croatia. Presumed synonym of A.
lendenfeldii (cf. Verhoeff, 1896). Syn.n.
Lysiopetalum cognatum. – Latzel 1888: 92. – Gasperini 1892:
š
8. – Karlinski 1894: 691. – Kovacević
1931: 71. – Stagl &
Stoev 2005: 8. – Stagl 2006: 227.
Lysiopetalum (Apfelbeckia) Lendenfeldii Verhoeff, 1896:
466, figs 1–4. Type locality: Bilek Cave in Bosnia and Herzegovina. Syn. n.
Lysiopetalum (Apfelbeckia) Lendenfeldii. – Verhoeff 1897:
š
154; 1899: 761. – Kovacević
1931: 71.
Apfelbeckia Lendenfeldii. – Verhoeff 1901: 275, figs 3, 6.
Apfelbeckia Enderleinii Verhoeff, 1901: 275, figs 1, 4. Type
locality: “Grabovica Höhle, Radoboljathal bei Mostar, Buchenwald am Prenj”, Bosnia and Herzegovina. Syn.n.
Apfelbeckia albosignata Verhoeff, 1901: 276, figs 2, 5. Type
locality: Schuma, Wolfshöhle, Bosnia and Herzegovina.
Syn.n.
Apfelbeckia silvivaga Verhoeff, 1901: 277. Type locality:
Jablanica, Bosnia and Herzegovina. Syn.n.
Apfelbeckia wohlberedti Verhoeff, 1909a: 717, 720, figs 1–6.
š
Type locality: Taubenhöhle (Tauben Cave) near Reci,
Albania. Syn.n.
Apfelbeckia silvivagum [sic!]. – Verhoeff 1909a: 720.
Apfelbeckia enderleini [sic!]. – Verhoeff 1909a: 720; 1943:
167. – Attems 1929: 322 (in the table, p. 291, spelled
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THE MILLIPEDE TRIBE APFELBECKIINI
A
B
D
C
E
F
Fig. 4. Apfelbeckia insculpta. – A–B. Syntype of Lysiopetalum insculptum L. Koch, 1867, BMNH, drawings by R. Hoffman. A.
Gonopod, mesal view. B. Same, posterior view. – C. Syntype of Lysiopetalum lendenfeldii Verhoeff, 1896, ZMB. Gonopod,
mesal view. – D. Syntype of Apfelbeckia duplocalca Attems, 1951, NHMW. Gonopod, posterior view. – E. Syntype of A.
lendenfeldi miraculosa Attems, 1951, NHMW. Gonopod, lateral view. – F. Syntype? of Karlabsolonia mirabilis Attems, 1951,
NHMW. Gonopod, posterior view. – b, coxal process 2; h, process on t; ib, inner branch; p, coxal process 1; ps, parasolenomere;
sg, seminal groove; t, tarsal process. – A–C, F without scale.
Stoev & Enghoff rettet.pmd
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P. STOEV & H. ENGHOFF
54
enderleinii); 1959: 303, 369, 373, figs 156–157. – Strasser
1971: 31. – Mršić 1994: 237.
Apfelbeckia albosignatum [sic!]. – Verhoeff 1909a: 720.
Apfelbeckia lendenfeldi [sic!] var. abbreviatum Verhoeff,
1909a: 720. Type locality: Jablanica, Bosnia & Herzegovina.
Synonymy proposed by Mršić (1994).
Apfelbeckia lendenfeldi [sic!] var. herzegowinense (spelled
herzegowinensis in figure caption) Verhoeff, 1909: 720. fig.
7. Type locality: unspecified site in North Herzegovina.
Synonymy proposed by Mršić (1994).
Apfelbeckia lendenfeldi [sic!]. – Verhoeff 1909: 721; 1943:
168, 170, figs 1–3. – Attems 1929: 322; 1959: 303, 369, 370,
fig. 146. – Strasser 1971: 31. – Mršić 1994: 237. – Mauriès et
al. 1997: 289.
Apfelbeckia wohlberedti. – Attems 1929: 322; 1959: 303,
369. – Verhoeff 1943: 168. – Strasser 1971: 31; 1976: 642. –
Moritz & Fischer 1974: 352. – Mauriès et al. 1997: 261, 289,
fig. 3C–E. – Makarov et al. 2004: 243. – Stagl & Stoev 2005:
20.
Apfelbeckia albosignata. – Attems 1929: 322; 1959: 303, 369,
375, figs 158–160. – Verhoeff 1943e: 167. – Strasser 1971:
31. – Mršić 1994: 237. – Mauriès et al. 1997: 289. – Makarov
et al. 2004: 243.
Apfelbeckia silvivaga. – Attems 1929: 322; 1959: 303. –
Verhoeff 1943: 166. – Tomić-Jovanović 1964: 193, 194. –
Strasser 1971: 31. – Moritz & Fischer 1974: 348.
Apfelbeckia hessei Verhoeff, 1929: 634, pl. VI, figs 35–40.
Type locality: “Ruinen von Salona bei Split”, Croatia. Syn.n.
Apfelbeckia lendenfeldii var. flavipes Attems, 1929: 322, 352.
Type locality: cave in Sildi Mountain near Shkodër (Scutari),
Albania. Syn.n.
Apfelbeckia lendenfeldi [sic!] var. abbreviata [sic!]. – Attems
1929: 322.
Apfelbeckia hessei. – Verhoeff 1932: 1507, figs 905–906;
1943: 167, figs 8–9. – Lang 1939: 291–293. – Attems 1959:
303, 369. – Strasser 1971: 31. – Mršić 1994: 237. – Mauriès et
al. 1997: 289.
Apfelbekia [sic!] lendenfeldi. – Lang 1935: 83–86.
Apfelbeckia albanica Verhoeff, 1941: 181, figs 1–6. Type
locality: Cukale Cave near Tirana, Albania. Syn.n.
Apfelbeckia albanica. – Verhoeff 1943: 168. – Attems 1959:
303, 369. – Strasser 1976: 642. – Mauriès et al. 1997: 289.
Apfelbeckia subterranea Verhoeff, 1943: 167, 169, figs 10–
12. Type locality: Novibazar District, Popova Cave near
Mileševo, 1040 m, Serbia. Syn.n.
Apfelbeckia lendenfeldii caligulifer Verhoeff, 1943: 167,
170, figs 4–5. Type locality: “Planjska-Höhle bei Lucovice
unweit Bilek”, Bosnia and Hercegovina. Synonymy proposed
by Strasser (1971).
Apfelbeckia lendenfeldii hebes Verhoeff, 1943: 167, 170, figs
6–7. Type locality: “Süd-Herzegovina, Grnkovaca-Höhle, ¾
Stunde entfernt von der Plavakirche b. Bilek”. Synonymy
proposed by Strasser (1971).
Apfelbeckia Hessei [sic!] var. Boldorii Manfredi, 1945: 6, figs
2–3. Type locality: Spela met Potzi, Kruja; Mali i Krujes; pr.
Spela met Potzi, Kruja, Albania. Syn.n.
Apfelbeckia duplocalca Attems, 1951: 256. Type locality:
Gorodnica Cave, Bosnia and Herzegovina. Syn.n.
Apfelbeckia lendenfeldi [sic!] miraculosa Attems, 1951: 256.
Type localities: Visocica and Mrcine in Bosnia and Herzegovina, and a cave at Dvršnik in Croatia. Syn.n.
ˆ
Stoev & Enghoff rettet.pmd
54
Karlabsolonia mirabilis Attems, 1951: 257. Type locality:
Jama (Cave) pod (below) Malim Kraljevcem, Croatia. Syn.n.
Apfelbeckia lendenfeldi miraculosa. – Attems 1959: 303, 369,
371, figs 147–155. – Strasser 1971: 31. – Mauriès et al. 1997:
289. – Guéorguiev et al. 2000: 229. – Stagl & Stoev 2005: 18.
Apfelbeckia duplocalca. – Attems 1959: 303, 369, figs 143–
145. – Strasser 1971: 31. – Stagl & Stoev 2005: 11.
Apfelbeckia subterranea. – Attems 1959: 303, 369. – Strasser
1971: 31. – Makarov et al. 2004: 243.
Karlabsolonia mirabilis. – Attems 1959: 303, 376, figs 161–
168.
Apfelbeckia Lendenfeldi [sic!]. – Tomić-Jovanović 1964: 193,
194.
Apfelbeckia mirabilis. – Strasser 1971: 9, 31. – Mršić 1994:
238. – Stagl & Stoev 2005: 18.
Apfelbeckia enderleinii. – Moritz & Fischer 1974: 335.
Lysiopetalum lendenfeldii. – Moritz & Fischer 1974: 339.
Apfelbeckia lendenfeldi var. herzegowinense. – Moritz &
Fischer 1974: 340.
Apfelbeckia Hessei [sic!] var. Boldorii. – Manfredi 1976: 224.
Apfelbeckia lendenfeldii flavipes [rank of subspecies, sic!]. –
Strasser 1976: 642. – Mauriès et al. 1997: 289.
Apfelbeckia hessei boldorii [sic!]. – Strasser 1976: 642.
Apfelbeckia lendenfeldii. – Makarov et al. 2004: 243. – Stagl
& Stoev 2005: 16.
Apfelbeckia lendenfeldii var. flavipes. – Stagl & Stoev 2005:
12.
Material examined:
Type specimens: Lysiopetalum insculptum: lectotype M,
paralectotype F, by present designation, L. Koch’s collection
Inv. No. 1913.7.25.418-19 [Dalmatien, Erber] (BMNH);
adult intact F (possible type), Inv. No. 4376, Acquisition
1866.I.53 [Dalmatia, coll. don. J. Erber 1861] (NHMW).
Apfelbeckia albosignata: M, broken into three pieces,
gonopods missing, syntype, Inv. No. A20051002 [Herzegowina, Schuma (Wolfshohle)] (ZSM). Apfelbeckia albanica: male gonopod broken into pieces, holotype, slide Inv.
No. A20034683 [Cukale Höhle, Albanien] (ZSM); 7 and 8
male leg-pair and antenna, holotype, slide No. A20034684
[Cukale Höhle, Albanien] (ZSM); 2 ad. FF broken into pieces,
[very likely non-types although labelled “syntype” by J.
Spelda, 4.iii.2005, but this species has been described from a
single male specimen], Inv. No. A20051001 [Albanien]
(ZSM). Apfelbeckia duplocalca: gonopods of 2 MM, adult F
broken into two pieces, syntypes, Inv. No. 2302 [Bosnia and
Herzegovina, Gorodnica Cave, coll. don. K. Absolon]
(NHMW). Apfelbeckia enderleinii: 1 adult F, 1 subad. F, 6
juv., syntypes, Inv. No. 2451 [Grabowica Höhle] (ZMB). 1 M
specimen broken into three pieces, gonopods missing; three
juveniles in another tube, syntypes, Inv. No. A20051007
[Herzegowina, Radoboljathal bei Mostar, labelled by J.
Spelda, 4.iii.2005] (ZSM). Apfelbeckia hessei: male gonopod broken into pieces, syntype, slide Inv. No. A20032682
[Salona] (ZSM); 6–9 male leg-pair, syntype, slide Inv. No.
A20032683 [Salona] (ZSM); 2–5 male leg-pair, syntype,
slide Inv. No. A20032684 [Salona] (ZSM); 2–5 male leg-pair
(second male), syntype, slide Inv. No. A20032685 [Salona]
(ZSM); one specimen broken into pieces (head, middle and
posterior PTs, telson), possibly male, gonopods missing, syntype, Inv. No. A20051003 [Croatia: ruinen von Salona bei
28-03-2008, 13:47
55
THE MILLIPEDE TRIBE APFELBECKIINI
Split, 3.iv.1928, leg. K.W. Verhoeff, labelled by J. Spelda,
4.iii.2005] (ZSM). Apfelbeckia lendenfeldii: gonopod, syntype, slides No. 827–828, Inv. No. 12923ab [Bilek-Höhle,
Bosnien, Apfelbeck leg.] (ZMB); adult M, 1 subad. M with
undeveloped gonopods, syntypes, Inv. No. 2115, Acquisition
1897.XXIII.2 [Bosnia, don. Verhoeff 1897] (NHMW). 4–7
male leg-pair, syntype, slide Inv. No. A20032686 [Bosnien]
(ZSM); 1–3 male leg-pair, syntype, slide Inv. No. A20032687
[Bosnien] (ZSM); head: antenna, gnathochilarium, labrum,
mandibles, syntype, slide Inv. No. A20032688 [Bosnien]
(ZSM); 1 specimen broken into pieces, possibly male, gonopods missing, syntype?, Inv. No. A20051010 [Herzegowina,
Bilek b. Pr. [?], labelled by J. Spelda, 4.iii.2005]; intact F,
syntype?, Inv. No. A20051012 [Bosnien, K.W. Verhoeff
Coll., labelled J. Spelda, 4.iii.2005] (ZSM). Apfelbeckia lendenfeldi caligulifer: gonopod broken into pieces, holotype,
slide Inv. No. A20034692 [Planjska-Höhle, K.W. Verhoeff
Coll.] (ZSM). Apfelbeckia lendenfeldii var. flavipes: 1 adult
M broken into pieces, gonopods in separate microtube, syntype, Inv. No. 4367 [Albania, cave in Sildi Mountain near
Shkodër (Scutari), leg. Petrovic
š 30.vii.1905, don. Petrovicš
1905] (NHMW). Apfelbeckia lendenfeldi genuina Verhoeff, 1943 [subsp. lendenfeldi]: 7–8 male leg-pair, slide
Inv. No. A20034685 [Mostar, Projecenica-Höhle,
K.W. Verš
hoeff Coll.] (ZSM); gonopod broken into pieces, slide Inv.
No. A20034686 [Mostar, Projecenica-Höhle,
K.W. Verhoeff
š
Coll.] (ZSM); several specimens broken into many pieces,
Inv. No. A20051014 [Herzegowina, in der Projecenica-Höhle
am Busak-Plateau bei Mostar, 680 m, 2.viii.1936, P. Remy
leg., labelled J. Spelda, 4.iii.2005]; gonopod broken into
pieces, slide Inv. No. A20034689 [Kali-Höhle, K.W. Verhoeff Coll.] (ZSM); 7–8 male leg-pair, slide Inv. No.
A20034690 [Kali-Höhle, K.W. Verhoeff Coll.] (ZSM).
Apfelbeckia lendenfeldi hebes: gonopod broken into pieces,
holo?type, slide Inv. No. A20034693 [Grnkovaca-Höhle,
š
K.W. Verhoeff Coll.] (ZSM). Apfelbeckia lendenfeldi var.
herzegowinense: gonopod, male leg-pair 8, syntype, slides
No. 3401–3402, Inv. No. 12940ab [Herzegowina, Verhoeff
leg.] (ZMB). middle and posterior segments of one broken
into pieces specimen, syntype, Inv. No. A20051004 [Drüsenloh[?], labelled by J. Spelda, 7.iii.2005] (ZSM). Apfelbeckia lendenfeldi miraculosa: 1 adult M with undissected
gonopods, gonopods in separate microtube, syntype, Inv. No.
2408 [Bosnia and Herzegovina, Visocica, coll. don. K. Absolon] (NHMW); 1 adult M and F, gonopods of second male,
syntypes, Inv. No. 4425 [Bosnia and Herzegovina, Mrcine,
4.viii.1912, coll. K. Absolon No: 87, don. K. Absolon]
(NHMW); numerous MM and FF, syntypes, Inv. No. 4426
[Bosnia and Herzegovina, Mrcine, 4.viii.1913, coll. K. Absolon No: 87, don. K. Absolon] (NHMW); 1 adult intact M,
syntype, Inv. No. 4427 [Dalmatia, Krivošije, upper cave near
Dvršnik (obere Höhle am Dvršnik), 6.viii.1912, coll. K. Absolon No: 668, don. K. Absolon] (NHMW). Apfelbeckia
silvivaga: gonopod, holotype, slide No. 1554, Inv. No. 12934
[Jablanica] (ZMB). male, broken into two pieces, gonopods
missing, holotype, Inv. No. A20051000 [Herzegowina,
Jablanica, labelled by J. Spelda, 3.iii.2005] (ZSM). Apfelbeckia subterranea: 7–8 male leg-pair, syntype, Inv. No.
A20034694 [Sandjak, Popova-Höhle] (ZSM); gonopod broken into pieces, syntype, Inv. No. A20034695 [Sandjak,
Popova- Höhle] (ZSM); M broken into pieces, gonopods
ˆ
Stoev & Enghoff rettet.pmd
55
missing, syntype, Inv. No. A20051005 [Herzegowina, im
Sandschak Novibazar, in der Popova-Höhle bei Mileševo,
1040 m hoch, 22.vii.1933, leg. P. Remy, labelled by J. Spelda,
4.iii.2005]. Apfelbeckia wohlberedti: 1 intact subad. M with
undeveloped gonopods, syntype, Inv. No. 4372 [Albania,
coll. Dr. Wohlberedt?, don. K.W. Verhoeff] (NHMW); 1
subad. M with undeveloped gonopods, syntype, Inv. No. 4535
[Albanien, Wohlberedt leg. Juni] (ZMB); 1 specimen broken
into pieces, possibly male, gonopods missing, syntype, Inv.
No. A20051006 [Albania, Taubenhöhle bei Reci in Albanien,
Juni, leg. O. Wohlberedt, labelled by J. Spelda, 4.iii.2005]
(ZSM). Karlabsolonia mirabilis: male gonopods in microtube; two slides of leg-pairs 1–4, 7–8, one female cyphopod
and “Gonopodentarsus”, syntypes, Inv. No. 2303 [Croatia,
pot-hole below Malim Kraljevica (Jama pod Malim Kraljevcem), Lok. 338, 29.ii.1914, coll. K. Absolon, don. K. Absolon] (NHMW).
Type specimens?: Lysiopetalum cognatum: slide with legpairs 7–9, Inv. No. 3990 (NHMW); adult intact F and one F
broken into pieces, 1 dissected M, gonopods in separate
microtube, Inv. No. 4382 [Dalmatia, Bosnia, don. R. Latzel
1919] (NHMW) (see also Stagl & Stoev 2005).
Old non-type material assigned to various nominal
(sub)species: Apfelbeckia enderleinii: two adult males
broken into pieces, gonopods missing, Inv. No. A20051015
[Herzegowina, Grabovica, 28.viii.1936, leg. P. Remy,
topotypes, J. Spelda det. 04.iii.2005] (ZSM); 1 juv., Cat. No.
3893 [Herzegovina, Verhoeff Coll.?] (AMNH); 1 juv., no
data, bought from Verhoeff (ZMUC). Apfelbeckia lendenfeldii: gonopod broken into pieces, slide Inv. No. A20034688
[Bilek, Kljenavacka[?]-Höhle,
K.W. Verhoeff Coll.] (ZSM);
š
gonopod broken into pieces, slide Inv. No. A20034691
[Herzegowina, Bilek-Höhle, Kuce[?]
Milovano[?], K.W.
š
Verhoeff Coll.] (ZSM); 3 FF broken into pieces, Inv. No.
20051008 [Herzegowina, K.W. Verhoeff det.]; M, broken
into pieces, gonopods missing, Inv. No. A20051009 [Herzegowina, Ramathalgrotte, Coll. K.W. Verhoeff, labelled J.
Spelda, 4.iii.2005]. 1 ad. F, Inv. No. 470 [Herzegovina,
Verhoeff Coll.?] (AMNH). Apfelbeckia lendenfeldi remyi
[undescribed taxon, preliminary name in collections]: gonopod broken into pieces, slide Inv. No. A20034687 [Herzegowina, Trebinje, Poganjaca-Höhle, K.W. Verhoeff Coll.]
(ZSM); one male broken into pieces, gonopods missing, Inv.
No. A20051013 [Poganjaca-Höhle, 27.viii.1936, K.W. Verhoeff Coll.]. Apfelbeckia wohlberedti: F, antennae, slide Inv.
No. A20034696 [Albanien [?], K.W. Verhoeff Coll.] (ZSM);
subad. M with undeveloped gonopods, Inv. No. 471 [Albanien Mts., Herzegovina, Verhoeff Coll.?] (AMNH); 1 F, 1
juv., Albania, Shkodër Distr., Daic, Spela Krovenices,
12.xi.1991, N. Lanjev leg., Golovatch, Mauriès & Stoev det.
(ZMUC).
New records: Bosnia and Herzegovina: 1 M, Provalija
pećina (pećina=cave), Kifino Selo, 23.vii.1967, P.R. &
C.L. Deeleman leg. (MHNG); 2 FF, juv., same locality,
8.viii.1968, H. Dekking, P.R. & C.L. Deeleman leg.
(MHNG); ad. M (48 PTs), ad. F (48 PTs), v. Kifino selo,
Snjetnica Cave, 20.vii.1962, P.R. & C.L. Deeleman leg.
(NNM). Croatia: 2 MM (one dissected, without gonopods,
second with 48 PTs). Zoölog. Mus. Amsterdam, Excursie
Jougoslavia, 1954, station 43, 16.v.1954, Dalmatià: Cavtat
(= Ragusa Vecchia) N42º35’, E18º12’, Sipun spilja (= Ae-
28-03-2008, 13:47
P. STOEV & H. ENGHOFF
56
sculapius Cave) (ZMAN); 1 F (48 PTs), Dalmatia: Cave
Mociljska spilja near village Pobrezje, near Dubrovnik,
17.vi.1961, Ellis leg. (ZMAN); 1 M (48 PTs) Dalmatia: Cave
Mociljska spilja near village Pobrezje, near Dubrovnik,
16.vi.1962, Exc. Zoöl. Mus. (ZMAN); 1 M, 2 FF, Dalmatia:
Cave Vilina Kuca
š in Ombla-dal, 16.vi.1962, Ellis leg.
(ZMAN); 5 MM, 8 FF (one ad. M and one ad. F each with 47
PTs; one ad. F 92 mm) Zoölog. Mus. Amsterdam, Excursie
Jougoslavia, 1954, station 41, Cave Vilina Pećina, cave at the
eastern end of the Ombla valley, 60 m depth? N42º40’,
E18º07’E, 16.v.1954 (ZMAN); 1 ad. M (48 PTs), cave near
Mocilje
near Dubrovnik, 28.v.1962, Yugoslavie
š (or Micelje)
š
exc. 1962 (NNM); 1 subad. M (47 PTs; undeveloped gonopods), same locality, 28.v.1962, J. Wiebes leg., Yugoslavie
exc. 1962 (NNM); 1 ad. M, 47 PTs, same locality, 28.v.1962,
exc. Leidse Biologen (NNM); M, F, island of Mljet,
vill. Ropa, cave Spilja pri Nereznom dolu, N42º45.512
E17º26.163, alt. 200 m, under stones, 13.viii.2006, B.
Petrov, S. Lazarov leg. (NMNHS). Montenegro: 1M, 1F,
Durmitor Mts., Tara River Gorge, Tmusha Cave, ca. 850 m
alt., hand coll., 17.vii.2003, S. Lazarov & M. Langourov leg.
(NMNHS); 2 MM, 1 F, Trnovo, Distr. Virpazar, cave Spilia,
alt. 360 m, N42º17’33.0’’ E19º02’10.0’’, under stones,
24.iii.2006, B. Petrov, S. Lazarov leg. (NMNHS); 1 F, Crni
nugli, Risan Distr., Dragalsko polje, Gorno Krivoshije,
Selakov dol, Chora pećina, alt. 750 m, N42º35’36.5’’
E18º42’42.1’’, under stones, 26.iii.2006, B. Petrov, S.
Lazarov leg. (NMNHS); 2 MM, 2 FF, 4 subad., vill. Seoce,
Distr. Virpazar, Golubova pećina, alt. 440 m, N42º12.509
E19º07.838, under stones, 12.viii.2006, t=14ºC, B. Petrov, S.
Lazarov leg. (NMNHS); 10 MM, 7 FF, 4 juv. (1 M & 1 F with
47 PTs), Zoölog. Mus. Amsterdam, Excursie Jougoslavia,
1954, station 51, valley of the river Lim near Andrijevica,
N42º47’, E19º48’E, 19–20.v.1954, deciduous forest (Corylus, Fagus) (ZMAN). Serbia: 2 MM, 1 F, Velika pećina,
Zvijezd, Prijepolje, 27.vii.1967, P.R. & C.L. Deeleman leg.
(MHNG); 2 MM, 1 F, Popova pećina, Sjelina, Prijepolje,
28.vii.1967, P.R. & C.L. Deeleman leg. (MHNG).
Diagnosis
Distinguished from congeners by the non-modified head in males in combination with the presence of two rows of pleurotergal setae, the absence of an apical, setiferous outgrowth on the
inner surface of the outer branch of the gonopod
femoroid which is terminally only slightly incised.
Distribution
The species distribution covers parts of Albania
(north and central parts south to Tirana), Montenegro, Bosnia and Herzegovina (central and
southern parts), Serbia (extreme west), Croatia
(southern parts of Dalmatia and island of Mljet).
Given the overall poor quality of Lang’s taxonomic work on millipedes (see, e.g., Enghoff
Stoev & Enghoff rettet.pmd
56
2006), the records of Apfelbeckia hessei from the
Croatian islands of Brach, Hvar, Korchula and
Vis (Lang 1939) should be treated with doubt.
Some of them, like those from caves near the
š on Brach, as
villages of Humac and Nerezišce
well as those from Hvar, are probably erroneous,
being most likely referable to A. brazzana.
Remarks
The most recent comprehensive work on Apfelbeckia so far is that of Attems (1959), where,
along with some re-descriptions, the author provided a key to the then recognized species belonging to the nominal subgenus, and having a
broader sponge-like process (k), thus excluding
wohlberedti, albanica and subterranea. Two
other “species”, namely A. silvivaga and A.
mirabilis, were not included in the key either.
Attems regarded Verhoeff’s subdivision as unsatisfactory because he used the length of the
antennae as the primary classification criterion.
Strasser (1962) first concluded that some of the
characters used for species separation are not
reliable and that the genus badly needs re-evaluation. He also mentioned that the angle of view
of the illustrations of gonopods has never been
taken into account by Attems, while in Verhoeff
(1943) it was indicated wrongly, and also expressed doubts about the precision of their drawings made from fragmented gonopodal parts
mounted on slides.
Having studied material from almost the
whole range of Apfelbeckia, we can conclude that
the importance of the shape of the sponge-like
process k has been overestimated by Attems.
This feature seems to characterize only different
populations and, taken separately, has no value
for species distinction. We did not find any clear
clinal variation in its shape, although it seems that
more southern specimens (e.g., those from Albania and Montenegro) usually have larger and
more elongated k (Fig. 4F), while smaller and
more equally ovoid k (like that of “A. enderleinii”) exist in specimens from the region of
Mostar, Bosna and Herzegovina, which forms
the north-northwestern border of the species’
distribution (Fig. 1). This and some other gonopodal characters, like the shape of the gonotarsus
and the setation on the inner and outer surface of
28-03-2008, 13:47
57
THE MILLIPEDE TRIBE APFELBECKIINI
A
B
C
D
Fig. 5. Apfelbeckia insculpta. Shape of gonopod tarsus in various nominal species, without scale. A. “enderleini”, non-type,
ZMB. B. “albosignata”, non-type, ZMB. C. “lendenfeldii”, non-type, ZMB. D. “duplocalca” syntype, NHMW.
the gonopods, do seem to vary between populations and cannot be regarded as species-specific. The colour and length of the antennae that
were used by Verhoeff (e.g. 1909a, 1943) cannot
provide enough support for clear separation of
the taxa that are here synonymized under A.
insculpta. Moreover, it is well known that coloration of specimens fades with the time of preservation of material. Although we cannot entirely
exclude the possibility of some of the taxa which
are here synonymised with A. insculpta being
valid, the available evidence in our opinion allows recognition of neither several species nor
even subspecies in this complex.
A similar situation concerns Acantopetalum
(Petalysium) carinatum (Brandt, 1840), another
callipodidan millipede confined to the western
part of the Balkan Peninsula and showing a distribution pattern similar to that of Apfelbeckiini. In
this species, the shape of the gonopodal so-called
U-process shows significant inter-populational
variability, also without any clinal pattern. Six of
the nominal (sub-)species forming the subgenus
Petalysium Strasser, 1976 were thus recently synonymised by Mauriès et al. (1997).
Lysiopetalum insculptum was described by L.
Koch (1867) from specimens collected in Dalmatia and Montenegro by Josef Erber. This species
remained in oblivion for a long time and its
taxonomic position has not been re-evaluated
since its original description. The sketches of the
gonopods made by Dr. R. Hoffman from the type
specimens in the BMNH (Fig. 4D, E) and the
examination of the female (type?) specimen in
Stoev & Enghoff rettet.pmd
57
the NHMW (see Stagl & Stoev 2005) leave
no doubt that this species is conspecific with
Apfelbeckia lendenfeldii. The latter, described by
Verhoeff in 1896, is the type species of Apfelbeckia and, after studying syntype specimens
from ZMB, we introduce here the new combination and synonymy: Apfelbeckia insculpta (L.
Koch, 1867) comb. n. = Apfelbeckia lendenfeldii
(Verhoeff, 1896) syn.n. Another possible synonym of Apfelbeckia insculpta is Lysiopetalum
cognatum Latzel, 1884 (cf. Verhoeff 1896),
described from a single female collected near
Dobrovnik (= Ragusa), Croatia.
There are neither gonopodal nor somatic characters, on the basis of which the nominal taxa
Apfelbeckia albanica, A. albosignata, A. duplocalca, A.enderleinii, A. hessei, A. hessei var.
boldorii, A. lendenfeldii var. flavipes, A. lendenfeldi miraculosa, A. subterranea and A. wohlberedti could be distinguished from A. insculpta.
So, having studied the respective type specimens, we propose here their synonomy under the
latter.
More complicated is the case of A. silvivaga,
the type species of the subgenus Haplobeckia,
whose holotype is in poor condition (a very
clumsy slide preparation of the gonopods, currently dried out and blackened) and cannot provide additional information about the status of
the species. Described from a single specimen
coming from the region where A. insculpta is also
known to occur (cf. Verhoeff 1909a sub A. lendenfeldtii abbreviatum), A. silvivaga can be presumed as having been based on a not fully mature
28-03-2008, 13:47
P. STOEV & H. ENGHOFF
58
male specimen with incompletely developed gonopods (without process k). Only further collecting from the type locality could answer this question, but for purely practical reasons we formalize here the synonymy with A. insculpta.
Examination of the type of Karlabsolonia mirabilis revealed a morphological aberration. In
the introduction of the catalogue of millipedes of
Yugoslavia, Strasser (1971) wrote that Hoffman,
after checking the type specimen in the NHMW,
had informed him in a letter that the presumable
penis of the species (figs 165–166 in Attems
1959) was actually the female ovipositor which
had inadvertently been mounted on the slide
together with male parts. We have also examined
the type material and can confirm that the slide
comprises a female cyphopod (“ovipositor”) together with the male gonotarsus. The remaining
part of the gonopods, preserved in alcohol, consist of a not fully developed, somewhat shrunk
and modified half, and a rudimentary second half
(Fig. 4E). Neither the brief original description
(Attems 1951) nor the subsequent, more detailed
one supplied with illustrations (Attems 1959),
which was compiled and published by Hans
Strouhal nearly 7 years after Attems’ death, has
any indication of the number and sex of the type
specimens. Having strongly malformed gono-
pods, as well as cyphopods at the same time, it is
very likely that Attems actually studied only one
specimen sharing both female and male sexual
characters, thus having mistaken the cyphopods
for a penis. This could be the first case of gynandromorphism in the Callipodida (see Discussion). Since currently there is not enough evidence to support a separate status of K. mirabilis,
we formalize its synonymy under A. insculpta.
Apfelbeckia synthesys sp. n.
Figs 1, 2, 6, 7, 9C
Material examined:
Holotype: adult male; 49 PTs, length 52 mm, width of midbody PT 3.23 mm; Greece, Olympos Mt., 700-2,100 m, 21–
26.v.1990, Zool. Mus. Copenh. Exp. (ZMUC) – Paratypes: 1
ad. male, 4 ad. females, 1 juvenile, same date, locality and
collector (ZMUC, ad. male & one female in the NMNHS). –
Non-type material: 1 F, 50 PTs, Greece, Macedonia, Olympos, 2100 m, Spilios Agapitos, 28.vii.–5.viii.1965, L.H.M.
Blommers leg. (ZMAN). There is a second label inside the
tube, “Acanthopetalum sp. det. Strasser, 1974”.
Etymology
The species is named after the Integrated Infrastructure Initiative SYNTHESYS (SYNTHEsis
of SYStematic resources), funded by the European Commissions’ 6th framework programme,
Table 1. Partial chaetotaxy in A. synthesys sp. n.
Anterior setae
Posterior setae
Collum
a, b, c, d, e, f, g, h+
a, b, c, d, e, f, g, h or
a, b, c, d, e, f, g+ a, b, c, d, e, f, g
-
2nd PT
a, b, c, d, e, f, g, h+
a, b, c, d, e, f, g, h or
a, b, c, d, e, f, g+ a, b, c, d, e, f, g
-
3rd PT
a, b, c, d, e, f, g, h+
a, b, c, d, e, f, g, h or
a, b, c, d, e, f, g+ a, b, c, d, e, f, g
-
4th PT
a, b, c, d, e, f, g, h+
a, b, c, d, e, f, g, h or
a, b, c, d, e, f, g+ a, b, c, d, e, f, g
-
5th PT
a, b, c, g + a, b, c, g
d, e, f, h + d, e, f, h
6th PT
a+a
b, c, d, e, f, g, h+ b, c, d, e, f, g, h
7th PT
a+a
b, c, d, e, f, g, h+ b, c, d, e, f, g, h
8th PT
usually all in posterior position
Stoev & Enghoff rettet.pmd
58
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59
THE MILLIPEDE TRIBE APFELBECKIINI
A
B
C
D
E
Fig. 6. Apfelbeckia synthesys sp. n. Male, holotype. A. Male gonopod, anterior view. B. Same, posterior view. C. Same, mesal
view. D, Same, anterio-mesal view. E. Same, posterio-lateral view. – a, coxal process 2; h, process on t; i, terminal incision of
femoroid; ib, inner branch; k, sponge-like process; n, broadened basal part of ib; ob, outer branch; p, coxal process 1; ps,
parasolenomere; s, solenomere; sg, seminal groove; st, sternum; t, tarsal process.
through the support of which the realization of
this project was made possible.
Diagnosis
Distinguished from congeners by the smaller
Stoev & Enghoff rettet.pmd
59
body size, the existence of only a single row of
pleurotergal setae and the following gonopodal
characters: apical part of femoroid strongly incised (vs. slightly or not incised) and process t
with a pointed hook-like process (h) (Fig. 6B, vs.
a more or less oval or notched process, Figs 3C,
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P. STOEV & H. ENGHOFF
60
A
B
C
Fig. 7. Apfelbeckia synthesys sp. n. – A–B. Female, paratype. A. Head and anterior PTs, lateral view. B. Cyphopods and second
leg-pair, anterior view. – C. Male, holotype. Leg-pair 6, anterior view. – B without scale.
4B). A single row of setae is also found in Himatiopetalum ictericum, but this taxon is well distinguished from the other apfelbeckiines by a
number of characters.
Description
Length: adult males: ca. 52 mm, adult females:
ca. 72 mm. Width of midbody PT: 3.2 mm
(males), 4.1 mm (females), 49–52 PTs + telson,
subadult male with 47 PTs.
Colour (Fig. 2): generally dark brownish; all
prozonites and posterior part of metazonites dark
brown, anterior part of metazonites light brownyellowish, more evenly yellowish around middorsal suture; lateral yellow spots present from
5th to ultimate pleurotergite, lying mostly on prozonite. Frons dark brown with four small, slightly
lighter spots near antennal bases and in the middle of the frontal part; stipes, cardo and epicranial
region marbled with yellow spots on dark brown
Stoev & Enghoff rettet.pmd
60
background; labrum chestnut, the zone above it
lighter. Antennae and legs dark brown, telson
chestnut.
Frontal part of head convex in both sexes,
densely covered with moderately dense, dark
setae. Ocellaria subtriangular, composed of ca.
35–40 (36 in holotype) black ocelli in 6–7 rows.
Tömösváry organs about three times larger than
an ocellus, placed between anterior line of ocellar
triangle and antennal base, well separated from
both. Antennae relatively short, extending to the
mid of PT 5 when folded backward (Fig. 7A).
All pleurotergal crests moderately developed;
crests on anterior PTs (excl. collum) flattened,
getting more pronounced caudad. Eighteen
crests between ozopores on PT 7. Ozopores usually on the 8th crest, placed at posterodorsal corner of a yellow spot. Chaetotaxy: see Table 1.
First and second leg-pairs markedly shorter,
third slightly shorter than subsequent legs. Tarsi
of leg-pairs 1–3 single; bi-articulated from 4th to
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THE MILLIPEDE TRIBE APFELBECKIINI
ultimate pair. All legs ending with a long and
curved claw. Coxal sacs present on legs 3–16.
Coxae of leg-pair 2 with posterior gonopores.
Coxa 7 short, subquadrate, without any particular
modifications; tibiae and tarsi in males with
stout, whitish pads (Fig. 9C). Tarsal pads larger
on anterior legs, getting smaller caudad. Hypoproct tripartite, medial sclerite largest, trapezoidal, bearing two paramedian macrosetae situated
close to each other; lateral sclerites with a seta
each. Paraprocts divided into small dorsal and
larger ventral sclerites, ventral sclerites bearing
ca. 14–20 scattered macrosetae along free borders. Spinnerets thin and short, ending with a
long macroseta.
Male gonopods (Fig. 6): in situ extending well
beyond pleurotergal edge, telopodites in touch
with each other. Sternum (st) – chitinized, subtrapezoidal, medially slightly incised, lying between 7th leg pair and telopodites. Coxa with two
processes – a large, apically rounded process (p)
covering the base of femoroid and about 1/3 of its
length, and a smaller, somewhat broadened and
apically slightly serrated one (a). Femoroid
strongly enlarged, divided into a large outer
branch (ob) and an inner branch (ib) ending with
a solenomere (s). Outer branch terminally incised
(i), bearing a sponge-like process (k), a large
anteromesal process (= tarsus, t), and an inner
hook-like process (h) pointing towards ib. Lateral side of outer branch densely covered with
macrosetae. Sponge-like process ovoid, placed
on posterior side of femoroid at about its
midlength, attached horizontally to it, and covered with stout dark spines contrasting against a
lighter background. Tarsus darkly coloured,
pointing downwards. Inner branch emerging at
about ¼ of femoroidal length, basal part strongly
broadened, forming a process (n). Inner branch
bifid; parasolenomere (ps) as long as solenomere
(s), both pointing towards inner surface of ob.
Seminal grove (sg) ending in upper branch.
Females: clearly larger than males, PTs 2 and
3 normal. Second leg-pair unmodified (Fig. 7B).
Remarks
Having only females and/or juveniles at our disposal, the above material could not be identified
closer to species.
Genus Himatiopetalum Verhoeff, 1900
Himatiopetalum Verhoeff, 1900a: 50. Type species: Lysiopetalum ictericum L. Koch, 1867, by monotypy.
Diagnosis: A genus of Apfelbeckiini with crested collum, undivided hypoproct, and outer femoroidal branch of gonopods flexible.
Himatiopetalum ictericum (L. Koch, 1867)
Figs 8, 9D
Lysiopetalum ictericum L. Koch, 1867: 895. Type locality:
Corfu, Greece.
Himatiopetalum ictericum. – Verhoeff 1900a: 50, 54, pl. 7,
figs 1–7; 1900b: 190; 1932: 1510, fig. 907, 1512, fig. 910. –
Attems 1926: 178, figs 215–216. – Strasser 1970: 241; 1974:
224, 289; 1976: 634. – Mauriès et al. 1997: 289.
Material examined:
Type specimens: 1 male lectotype, 1 male, two females and
one specimen of unknown sex, paralectotypes, by present
designation, Inv. No. 1913.7.25.713-717 [Corfu, Erber 346]
(BMNH; R. Hoffman & J. Beccaloni, in litt.).
Other specimens: entire undissected M [Corfu, K.W. Verhoeff Coll.] (ZMB); slide of gonopods [Corfu, K.W. Verhoeff
Coll.] (ZMB); 1 F, Inv. No. 472 [Corfu, Verhoeff Coll.?]
(AMNH); ad. M [Corfu] (NHMW); 1 F, no data, bought from
Verhoeff (ZMUC).
Distribution
Apfelbeckia sp.
Material examined:
Albania: 1 F, Mt. Gaitan, v. Konak de Gaitan, Shkodër,
Stoev & Enghoff rettet.pmd
10.xi.1991, N. Landjev leg. (NMNHS). Bosnia and Herzegovina: 1 M, Ledenica pećina, Kalinovik, 3.viii.1968, P.R. &
C.L. Deeleman leg. (MHNG). Croatia: 1 M (dissected, without gonopods), Zoölog. Mus. Amsterdam, Excursie Jougoslavia, 1954, station 33, 12.v.1954, Dalmatia: Malapetsj near
Sutina, in the Mosor Planina Mt., S of Sinj; small cave, about
50 m long, stalactites, collapsed doline (ZMAN); 1 juv.,
Dalmatia, Cave Vranjac near village Kotlenice, near Dugopolje, 7.vi.1961, W.N. Ellis leg. (ZMAN). Montenegro: 3
FF, Grahovo, Distr. Niksic, Gorno Krivoshije, cave Dakovica
pećina, alt. 700 m, N42º39’22.4’’ E18º40’38.5’’, under
stones, 28.iii.2006, B. Petrov, S. Lazarov leg. (NMNHS).
Serbia: subad. M, juv., Gornja pećina kod Petnice, Valjevo,
9.vii.1976, P.R. & C.L. Deeleman leg. (MHNG).eci
61
Known only from the island of Corfu, Greece.
(There is, however, material in the NHMW collected from the Llogorasë Pass, south of the city
28-03-2008, 13:47
P. STOEV & H. ENGHOFF
62
A
C
B
D
Fig. 8. Himatiopetalum ictericum. Non-type specimens, ZMB, NHMW. A. Head and anterior PTs. B–C. Gonopod, mesal view.
D. Gonopod, anterior view. – cp, coxal process; ib, inner branch; k, sponge-like process; m, process on the inner surface of ob;
sg, seminal groove; ob, outer branch; st, sternum.
of Vlorë, Albania, putatively referred to as H.
ictericum by Attems.)
Remarks
Since all existing illustrations of the species
come from earlier works of Verhoeff, and for
consistency with the descriptions of other species
of Apfelbeckiini, here we provide new illustrations of the head and anterior PTs, male leg-pair
7 and gonopods.
Key to the species of Apfelbeckiini
1. Outer femoroidal branch (ob) of gonopods
Stoev & Enghoff rettet.pmd
62
flexible, entirely enveloping the inner branch;
coxal process (cp) membranous, enveloping the
femoroidal base; sponge-like process (k) with
elongated spines, spines not tapering, equally
broad along their length; tarsal process (t) missing; (Fig. 8B–D); trochanter of male leg-pair 7
expanded dorsally (Fig. 9D); hypoproct undivided; collum with well pronounced crests (Fig.
8A); midbody PTs with crests below the level of
ozopores ……........... Himatiopetalum ictericum
– ob solid, partly enveloping the inner branch;
coxal process (p) hard, not membranous, placed
laterally; sponge-like process with short tapering
spines; tarsal process (t) present; trochanter of
male leg-pair 7 normal (Fig. 9A–C); hypoproct
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THE MILLIPEDE TRIBE APFELBECKIINI
A
B
C
D
Fig. 9. Male leg-pair 7, anterior view, without scale. A. A. brazzana, paralectotype, NHMW. B. A. insculpta, non-type, NMNHS.
C. A. synthesys, holotype, ZMUC. D. H. ictericum, non-type, ZMB.
tripartite; collum smooth; midbody PTs without
crests bellow the level of ozopores …..............
Genus Apfelbeckia ........................................... 2
2. Male head concave frontally; inner surface of
ob with an apical outgrowth bearing thick setae
(g in Fig. 3B) ……………............... A. brazzana
– Male head convex; inner surface of ob without
outgrowth …..……...........................………… 3
3. Terminal part of ob strongly incised; hook-like
process (h) pointed; sponge-like process (k)
small; inner side of ob devoid of setae (Fig. 6);
PTs 1–4 with a single row of setae in anterior
position; small species (up to 50–60 mm); antennae in males short, reaching the mid of PT 5 when
folded backward ……….................. A. synthesys
Stoev & Enghoff rettet.pmd
63
– Terminal part of ob slightly incised; h rounded
or notched; k larger; inner side of ob with setae
(Fig. 4A–D); PTs 1–4 with two rows of setae
in anterior and posterior position; larger species
(up to 100 mm); antennae in males long, extending beyond PT 7 when folded backward ..............
......................................................... A. insculpta
DISCUSSION
In their paper on Turkish Callipodida, which
constitutes the starting point of modern callipodidan taxonomy, Hoffman & Lohmander (1964)
remarked about the family Schizopetalidae that
“Species in this family are moderate to fairly
28-03-2008, 13:47
P. STOEV & H. ENGHOFF
64
large in size, and offer good materials for the
study of geographic dispersal and variation. A
beginning of such work is herewith made, in the
form of a preliminary revision of Eurygyrus and
it is to be hoped that someday similar studies can
be made on such large genera as Acanthopetalum
and Apfelbeckia”. The present paper is a partial
fulfillment of Hoffman & Lohmander’s hope,
although the outcome of our work on Apfelbeckia
et al. is quite different from that of theirs on
Eurygyrus. Whereas the latter genus is still regarded as being composed of a large number of
species (18, according to the most recent contribution (Stoev 2007)), we have synonymized all
previously described species of Apfelbeckia under one name. In this respect, our study is more in
line with the recent detailed studies on the genus
Rhymogona Cook, 1896 (Chordeumatida: Craspedosomatidae), in which previous authors (notably Verhoeff) had described a host of taxa
which, on closer scrutiny, turned out to fall within a single biological species (Pedroli-Christen &
Scholl 1996, but see Spelda 2005 for a contrasting view). Without doubt, revision of several
other genera of European millipedes will result in
similar massive synonymisations.
Of the three species we recognize in Apfelbeckia, gonopodal similarity is particularly
strong between the insular species A. brazzana
and the mainland species A. insculpta, suggesting recent separation of the island populations
from the mainland ones. We are, however, unable to present a proper phylogenetic analysis of
relationships between apfelbeckiines, the available evidence being too meagre for this.
One specimen (type specimen of Karlabsolina mirabilis) seems to be the first example of
gynandromorphism in Callipodida. Gynandromorphism in millipedes has so far been docu-
mented only in species of the order Julida (e.g.,
Bigler 1920; Enghoff 1985).
ACKNOWLEDGMENTS
We thank J. Dunlop (ZMB), V. Stagl (NHMW),
P. Schwendinger (MHNG), H. de Jong and B.
Brugge (ZMAN), E. van Nieukerken and R. de
Vries (NNM Leiden), L. Prendini (AMNH), J.
Spelda and S. Friedrich (ZSM), and J. Beccaloni
(BMNH) for arranging the loan of type specimens or giving information about and access to
the respective collections under their care. We
are very grateful to R. L. Hoffman for sharing
with us his unpublished observations of the type
of Lysiopetalum insculptum and also for making
the sketches of the gonopods available for this
study. Mrs. Fani Bozarova, technician in the
NMNHS, inked part of the illustrations, G. Brovad (ZMUC) took the photograph of the new
species. The travels of the first author to different
European and American museums and the work
there was supported by the European Commission’s programmes “Transnational Access to
Major Research Infrastructures” (COBICE),
and from the SYNTHESYS Project (AT-TAF,
DK-TAF, NL-TAF) http://www.synthesys.info/
which is financed by the European Community
Research Infrastructure Action under the FP6
“Structuring the European Research Area” Programme, the National Science Foundation PEET
grant, the Field Museum Bass Scholarship, as
well as by Deutsche Forschungsgemeinschaft.
P.S. thanks B. Petrov (NMNHS) and S. Lazarov
(Institute of Zoology, Sofia) for contributing to
this study by collecting fresh material of A.
insculpta and A. brazzana in Montenegro and
Croatia in 2006.
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