Dorcadion pusillum pusillum Küster, 1847

Dascălu, Maria-Magdalena, 2018, The subspecific structure of Dorcadion (Cribridorcadion) pusillum Küster (Coleoptera: Cerambycidae) with description of two new subspecies from Romania, Zootaxa 4442 (1), pp. 43-62: 44-50

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Dorcadion pusillum pusillum Küster, 1847


Dorcadion pusillum pusillum Küster, 1847 

Figs 1A–D, F–GView FIGURE 1, 3View FIGURE 3, 5A–EView FIGURE 5, 6CView FIGURE 6.

Dorcadion pusillum Küster, 1847: 90  (original description). Syntypes ♂ ♀, not examined (Museum of Natural History Nürnberg?). Type locality: Podolia, South Russia, Greece.

D. pusillum v. berladense Pic, 1903: 7  (original description). Lectotype ♂, present designation ( MNHNAbout MNHN). Type locality: Moldavie, Val du Berlad [ Romania, Zorleni village according to Montandon (1908)].

D. pusillum  m. postdisjunctum Breuning, 1946: 100 (unavailable name). Type locality: Romania, Vallée du Berlad [ Zorleni village ]. 

Dorcadion pusillum  ; Kraatz, 1873: 75 (redescription).— Ganglbauer, 1884: 450 (keyed).— Montandon, 1908: 100 (faunistic list).— Plavilstshikov, 1958: 149 –151 (redescription, illustrated).— Panin & Săvulescu, 1961: 389 (redescription, distribution).— Breuning, 1962: 112, 168, 177, 286, 287 (keyed, redescription).— Kasatkin, 2006: 91 (endophallus structure).— Lazarev, 2009: 198.— Danilevsky, 2010: 251 (catalogued).— Lazarev (2011): 257.

Dorcadion pusillum pusillum  ; Kasatkin, 2002: 277 –280 (illustrated, distribution).— Danilevsky, 2010: 251.

D. pusillum v. berladense  ; Montandon, 1908: 100.

D. pusillum  ab. berladense  ; Plavilstshikov, 1958: 149, 150.— Panin & Săvulescu, 1961: 389.

D. pusillum  ab. postdisjunctum; Plavilstshikov, 1958: 149, 150.

D. pusillum  ab. postdijuncta; Panin & Săvulescu, 1961: 389.

D. pusillum  m. berladense  ; Breuning, 1962: 112, 288.

D. pusillum  m. postdisjunctum; Breuning, 1962: 159, 160, 287.

D. pusillum berladense  ; Kasatkin, 2002: 277 (change of status). — Danilevsky, 2010: 251 (catalogued).

Description. Male. Body length: 7.4–12.1 mm (n = 95), body width: 2.8–4.6 mm (n = 86). Body size varies between studied populations, with specimens from northern populations tending to be larger ( Table 1). Body black. Antenna uniformly dark brown to reddish-brown ( Fig. 3AView FIGURE 3) and first joint sometimes lighter. Legs black to dark brown, but femora and tibiae variably extensively reddish basally or fore tibia entirely reddish. Pronotum 0.78– 1.11× as wide basally as long (n = 86), with setiferous punctures distinctly separated mesally and denser and partly conjugated on sides; lateral tubercle small, with acute, moderately long spine. Pronotum dorsally with brown background pubescence, a median white stripe and sparse whitish pubescence laterally, mostly on tubercle. Elytra moderately convex, with dense, fine microsculpture and sparse small setiferous punctures; punctuation deeper and denser on humeral depression. Humeral carina roughly sculptured basally and well visible in basal third of elytra. Background elytral pubescence blackish-brown; humeral and dorsal stripes rusty or light brown, sometimes mixed with grayish pubescence towards apex ( Fig. 3B, C, H, JView FIGURE 3), well-developed but not strongly contrasting with background pubescence; usually stripes fused at elytral apex. Spots of velvety black pubescence present on dorsal and humeral stripes and along sutural stripe, forming a mottled elytral pattern ( Fig. 3D, F, H, J, K, PView FIGURE 3); pubescence of these spots caducous, in older specimens dorsal and humeral stripes interrupted by black elytral chitin ( Fig. 3G, I, L, NView FIGURE 3). Sometimes humeral and dorsal stripes inconspicuous, dark-brown, with reduced black spots of velvety pubescence ( Fig. 3E, M, OView FIGURE 3). Sutural stripe and a small humeral spot with white pubescence. Small, more or less confluent spots of velvety black pubescence form an interrupted subsutural stripe along sutural stripe. Lateral stripe white on epipleuron and mottled above it, with off-white to rusty pubescence interrupted by background pubescence.

Endophallus. Basal tube with small and weakly sclerotized ventral plates; medial tube slightly curved ventrally and enlarged distally forming a central bladder with distinct ventral and dorsal swellings; region before central bladder slightly swollen forming a second smaller bladder; central trunk covered in microspicules and connected at a wide angle to central bladder resulting in a distinct central bend; preapical bulb spherical and delimited from central trunk through a constriction; apical bulb spherical or more or less cylindrical (when not fully inflated) and covered in microspines ( Fig. 6CView FIGURE 6).

Female. Body length 9–13.9 mm (n = 42). Body size varies between studied populations, with specimens from northern populations tending to be larger. Females are commonly autochrome, and besides the biometric traits related to sexual dimorphism, differ from males in the colour of the stripes and of the background pubescence on the pronotum and elytra. Pronotum transverse with comparatively longer lateral spines. Elytra wider and more convex, with dense, brown to light brown background pubescence ( Fig. 5A, B, D, EView FIGURE 5) and with more developed humeral carina. Humeral stripe usually wider than dorsal (about as wide as interval between humeral and dorsal stripes or wider), stripes almost always fused at elytral apex. Because of lighter coloured elytral stripes (with light brown, creamy or off-white pubescence, sometimes mixed with rusty pubescence) the spots of velvety black pubescence on dorsal and humeral stripes are more contrasting than in males ( Fig. 5B, DView FIGURE 5); rarely, these spots are reduced in size and number ( Fig. 5A, EView FIGURE 5). Sutural stripe creamy to off-white. Lateral stripe as in male but wider.

Very few females are androchrome ( Fig. 5CView FIGURE 5), with dark background pubescence and rusty elytral stripes. In some populations (Hălceni, Galaţi) the autochrome and androchrome colour pattern intergrades so the separation between the two categories of females becomes arbitrary.

Remarks. Dorcadion pusillum  was described by Küster (1847) without a precise type locality from three distinct regions. The syntypic material of D. pusillum  contains specimens from Podolia (the present-day Ukrainian Vinnytsia and Khmelnytskyi areas), South Russia and Greece. No other record of D. pusillum  from Greece is known so far apart from Küster’s description. The reason of the inclusion of Greece will be later clarified by Kraatz (1873) according to whom the “griechische apicale Wattl in litt.”, placed under synonymy with D. pusillum  by Kuster (1847), is a synonym of D. minutum Kraatz, a species from Greece. Ganglbauer (1884) considered apicale as distinct and described it as D. minutum var. brenskei Ganglbauer  (currently accepted as D. brenskei according to Pesarini & Sabbadini 2004). Certainly Greece was originally included in the distribution of D. pusillum  based on a species from the minutum species group, as Kraatz (1873) has already shown.

According to Tavakilian & Chevillotte (2016) the syntypic material of D. pusillum  is deposited in the Museum of Natural History in Nürnberg. Unfortunately, it was not possible to obtain any further information and eventually designate a lectotype. As South Russia is a large region and also the type locality for the recently described D. pusillum tanaiticum  , designation of Podolia as type locality is preferable. Podolia is a large historical region situated in Ukraine to the north of present day Republic of Moldova and for this reason, in the maps from Figs. 7View FIGURE 7 and 8View FIGURE 8 the type locality was placed in the forest steppe biome, somewhere between Khmelnytskyi and Vinnytsia, where Küster’s specimens are most likely to have originated.

D. pusillum v. berladense  . It was described by Pic (1903) from “ Moldavie: Val du Berlad” based on male specimens collected by A. L. Montandon, a French naturalist and entomologist who lived and collected in Romania for most of his life ( Andrei & Pandele 2006). In MNHNAbout MNHN the locality label and the original “ type ” label in Pic’s handwriting are on a female specimen ( Fig. 1BView FIGURE 1) that cannot be the type as Pic described v. berladense  base on the male sex. Even if Pic gives a single body length for his variety, the material is stated to be deposited in two collections (Nicolas and Pic). Hence the single male specimen bearing an identification label written by Pic ( Fig. 1AView FIGURE 1) should be considered as syntype. For the stability of nomenclature, I here designate this specimen as lectotype.

Pic’s variety was regarded by Plavilstshikov (1958) as an aberration and by Breuning (1962) as a morpha. Kasatkin (2002) suggested, probably based on the description and distribution (type locality situated in the western margin of the species range), that the variety described by Pic has to be regarded as a subspecies. All specimens of D. pusillum  collected by A. L. Montandon and found in the natural history museums in Paris and Bucharest ( MNHNAbout MNHN and MGABAbout MGAB) have the same locality label “ Moldavie, Val. du Berlad”—a large area that covers the territory along the 200 km long Bârlad River, in present day eastern Romania (historical region Moldova). However, in the catalogue of the Romanian coleopteran fauna published five years after the description of v. berladense, A. L. Montandon ( Montandon, 1908: 100)  mentioned D. pusillum var. berladense Pic  only from Zorleni, a village in Bârlad River valley. Even if the labels do not indicate the exact locality, Zorleni is by inference the type locality for v. berladense  . Some specimens of D. pusillum  from “Val. du Berlad” in both MNHNAbout MNHN and MGABAbout MGAB collections were historically identified as D. decipiens Germar  ( Fig. 1D, FView FIGURE 1). The record of D. decipiens  from Zorleni by Fleck (1906) based on specimens from A. L. Montandon was hence connected with D. p. pusillum  , as males of D. decipiens  are superficially similar to the former except body size and elytra shape.

The availability of v. berladense Pic, 1903  . Traditionally, the varieties described by Pic are considered available names in accordance with article 45.6.4 of ICZN (1999) which stipulates that the name following a binomen is subspecific if the author published it before 1961 expressly under the term variety. However, the same article stipulates that a variety is unavailable if “ the author expressly gave it infrasubspecific rank or the content of the work unambiguously reveals that the name was proposed for an infrasubspecific entity ” ( ICZN, 1999). In the same work Pic (1903) describes, besides numerous varieties, a new subspecies as well: “ Leptura bitlisiensis Chevr.  s. esp. armeniaca ” ( Pic, 1903: 4). He also describes Pogonocherus taygetanus  from Greece asking himself if the new taxon could be related to P. plasoni Ganglbauer  or even considered as a subspecies of the latter: “Ne connaissant pas Plasoni Gglb. en nature je ne puis me rendre compte exactement si taygetanus  peut être considéré comme espèce voisine, ou regardé plutôt comme sous-espèce…” ( Pic 1903: 8). He raises the same question for Leptura excisipes Daniel  , namely if it might be the race (or subspecies) of another Leptura  L. species: “Je me demande si L. excisipes  ne serait pas la race (ou sous-espèce) asiatique de pallens Brul?” ( Pic, 1903: 4).

Taking into account the article 45.6.4 of ICZN (1999) and the guidelines for its interpretation by Lingafelter & Nearns (2013), the fact that Pic (1903) described and discussed both subspecies and varieties within the same work, means that varieties could be treated as infrasubspecific and in this case berladense  should be considered an unavailable name. A similar interpretation of this article was made by Bezdĕk & Regalin (2015) who considered the varieties of Crysomelidae species described by Pic in several papers as infrasubspecific because these papers contain at least one description of subspecies and thus, “the author himself stated for these varieties a lower level than the subspecific one” ( Bezdĕk & Regalin 2015: 6).

However, it is not easy to understand if v. berladense  has a subspecific or infrasubspecific rank from its original publication. In a paragraph immediately below the one describing P. taygetanus, Pic (1903)  states that a specimen of Saperda perforata Pallas  from Algeria should be distinguished from European specimens as a geographical variety: “Je dois à l’extrême obligeance de M. Quittard un exemplaire ♀ de Saperda perforata Pall.  provenant de Philippeville, en Algérie, exemplaire qui, à mon avis, mérite d’être distingué de nos exemplaires européens, a titre de var. ( v. algerica  )” ( Pic 1903: 8).

Therefore, in the spirit of the law the term variety in Pic (1903) has at least in one instance a connotation of subspecies, which in general usage of taxonomy was a replacement for “variety“ in its meaning of geographic race ( Mayr 1970: 210) and in this case berladense  should be an available name. But in the letter of the law, because Pic (1903) uses the term “variety“ for geographical forms in parallel with the term subspecies (and hence for a lower level of differentiation than subspecies), the name “variety, var., v.” is infrasubspecific (below subspecies).

For berladense  , an accurate interpretation is hard to make and the conclusion depends on whether one favors the spirit or the letter of the code. As the work ( Pic 1903) doesn’t indicate unambiguously the infrasubspecific rank and Pic’s varieties are routinely accepted as available names, I consider that berladense  is available.

The taxonomic status of populations attributable to v. berladense  . Irrespective of the availability of berladense  , the next question is whether the population from “Vall. du Berlad” (Zorleni) can be considered as a valid subspecies. According to the original description ( Pic 1903), D. pusillum v. berladense  is characterized by a more uniform and extensive dark pubescence on elytrae and legs with the colour slightly obscured: “ pattes a coloration un peu obscurcie, élytres ornes d’une pubescence plus uniforme ou d’une pubescence foncée plus étendue ”. The short description given by Plavilstshikov for berladense  (the dorsal and humeral stripes and the elytral spots as well are inconspicuous on the background coloration of elytra) could have been based also on specimens of D. pusillum tanaiticum  as a specimen from Rostov on Don identified as aberration berladense  by Plavilstshikov was included in the type series of D. p. tanaiticum ( Kasatkin 2002)  .

The males from the same series as the lectotype of v. berladense  (preserved in MGABAbout MGAB and MNHNAbout MNHN) have wornout pubescence and only one of them corresponds with Pic’s description ( Fig. 1CView FIGURE 1); other five have evident elytral stripes, though they are partly abraded ( Fig. 1D, FView FIGURE 1) and one is too mouldy to accurately determine the pattern. Breuning (1946) describes from “Vallée de Berlad“ D. pusillum  m. postdisjunctum based on a male similar with the nominal subspecies (i. e. with dorsal and humeral stripes), except dorsal stripe not fused with the humeral. Moreover, Montandon (1908) records from Zorleni both D. pusillum  and D. p. v. berladense  which again suggests a mixed series of specimens from the same locality.

The specimens from Sălcioara (at 13 km from Zorleni, the type locality of v. berladense  ) have the habitus of the nominal subspecies ( Fig. 3L, NView FIGURE 3) but a few males are darker, with humeral and dorsal stripes inconspicuous ( Fig. 3M, OView FIGURE 3) as in the lectotype. Darker males are rarely found also in other populations of the nominal subspecies, e.g. in Republic of Moldova at Răzeni and Tomaiul Nou ( Fig. 3EView FIGURE 3). Furthermore, based on the male elytral pattern, the distribution range of the nominal form extends both north and south of Zorleni, at Hanu Conachi and Galaţi (localities 18 and 19 on the map in Fig. 7View FIGURE 7). Because of the above reasons, the populations from Zorleni area cannot be separated as a distinct subspecies.

The darker individuals corresponding with Pic’s description for v. berladense  represent intrapopulational variability and berladense  is treated here as a synonym of the nominal subspecies.

Distribution. Ukraine, Crimea, Republic of Moldova, Romania ( Fig. 7View FIGURE 7).

Material examined. REPAbout REP. OF MOLDOVA: Ungheni, Elizavetovca  , 16.V.2013, leg. Dascălu, Fusu & Chinan (13♂♂, 3♀♀); Raion Ialoveni, sat Răzeni [Ialoveni district, Răzeni village], 26.III.2010, leg. A Zubov (11♂♂, 7♀♀), 20.IV.2011, leg. Dascălu, Fusu & Zubov (10♂♂, 8♀♀); 20 km de Hâncești, între Cneazevca și Tomaiul Nou [20 km from Hâncești between Cneazevca and Tomaiul Nou ]  , 19.IV.2011, leg. Dascălu, Fusu & Zubov (19♂♂, 2♀♀) [ MDCO]. Chişinău  , 29.III–11.IV.1923 (1♀), 5.IV.1924 (1♀), 20.IV.1925 (2♂♂, 1♀); Durleşti [near Chişinău] V.1922 (1♂)  , IV.1923 (1♂), V.1925 (1♀); Dănceni [near Chişinău] 22.V.1932 (1♀) [NMENH]. ROMANIA: Călăraşi [Botoșani Co.], 13.IV.1972 (1♀) [ NSMD]  ; Iaşi, dig lac Hălceni, 2.V.2006, leg. Dascălu & Fusu (8♂♂) [ NSMG]  ; Iaşi, Vlădeni, dig lac Hălceni [dam of Hălceni lake ], 15.IV.2000 (1♀)  , 29.IV.2004 (1♂, 1♀), 25.V.2005 (1♂), 2.V.2006, (7♂♂, 4♀♀), 13.V.2006 (1♂), leg. Dascălu & Fusu, 3.V.2007, leg. Fusu L. (1♀); Iași, Vlădeni, vale Miletin [ Miletin valley ]  , 02.V.2011 (2♂♂), 20.V.2011 (8♂♂), leg. Dascălu & Fusu; Iași, Rezervaţia Valea lui David [Valea lui David Reserve], 17.IV.2011, leg. Dascălu, Fusu & Chinan (11♂♂, 6♀♀ autochrome, 1♀ androchrome)  , 18.IV.2008 (5♂♂), 27.IV.2011 (1♂, 1♀), leg. Dascălu & Fusu, 6.V.2008, leg. Fusu L. (2♂♂, 1♀) [ MDCO]; V. David [Iaşi, Valea lui David Reserve]  , 9.V.1967 (1♂) [NSMI]; Iași, Leţcani, 14.V.2006, leg. Fusu L. (2♂♂, 2♀♀) [ MDCO]  ; Iași, Breazu  , 5.V.1961, leg. A. Popescu Gorj (1♀); Breazu jud. Iași  , 20.V.1958 and 6. VI.1958, leg. A. Popescu-Gorj (2♀♀) [ MGABAbout MGAB, Nr. 206, Serafim coll.]; Iaşi  , 18.V.1955, leg. M. Ieniştea (2♂♂); Breazu, Iaşi  , 5.V.1962, leg. A. Popescu-Gorj (1♂) [ NSMG]; Mârzeşti [Iași, Rediu village , Mârzeşti meadows]  , 21.V.1961 (12♂♂, 5♀♀); P. Sadoveanu [Iaşi, Sadoveanu Park] 16.V.1958, [probably leg. C. Mândru] (1♀) [ NSMI]  ; Iaşi, 18.V.1955, Dr. N. Săvulescu (1♂) [ MFBI]  ; Vaslui, Sălcioara , 11.V.2008, leg. Dascălu M. M  . (6♂♂) and 28.IV.2011, leg. Dascălu, Fusu & Chinan (10♂♂, 3♀♀) [MDCO]; Galaţi, Grădina Botanică [ Botanical Garden ], 4.V.2007, leg. Patriche G  . (1♀ androchrome), 13.V.2011, leg. Cristescu M. (6♂♂, 2♀♀ autochrome, 3♀♀ androchrome); Galaţi, 150 m  , 4. IV.2005 (1♀ androchrome), 6. IV.2005 (2♂♂, 1♀), 9. IV.2005 (1♀ autochrome, 2♀♀ androchrome), leg. A. Ruicănescu; Galaţi, Hanu Conachi 17.V.2005, leg. A. Ruicănescu (1♂) [ MDCO]  ; Galaţi, 4. IV.2005 (2♂♂)  , 6. IV.2005 (6♂♂, 2♀♀), 9. IV.2005 (10♂♂, 3♀♀), 10. IV.2005 (3♂♂, 5♀♀), leg. A. Ruicănescu [ NSMG]. Galaţi: Galatsch / Museum Paris Coll. J. Thomson 1952 / Ex. Musaeo James Thomson / Galaţi ( Roumanie) M. Al. Ieniştea corr. (1♀ androchrome)  ; Galaţi: pusillum Kust. Galavsch  [?] / Galaţi ( Roumanie) M. Al. Ieniștea corr. / Museum Paris Coll. M. Pic. (1♂) [ MNHNAbout MNHN]  ; Tulcea, Dobrogea, R. Jeannel, IV.26 / Museum Paris, Coll. R. Jeannel 1931 (5♂♂ of which 1♂ with the identification label “ Dorcadion pusillum Kust., Breuning  det.”, 3♀♀ autochrome, 1♀ androchrome) [ MNHNAbout MNHN]  ; Galaţi (1♂, Fig. 1GView FIGURE 1): Galatsch [picture examined, Kraatz coll., SDEI]  . UKRAINE: Odessa Oblast, Ovidiopol, Liman Nistru Border, 2.V.2013, leg. Dascălu, Fusu & Fusu (10♂♂, 4♀♀)  ; Kherson region , Kherson city, 24.IV– 9.V.1996, leg. Mishustin R. (4♂♂) [ MDCO]  ; Odessa (1♂) [picture examined, Kraatz coll., SDEI]  ; Odessa (1♂): pusillum    , Odessa / Museum Paris Coll. J. Thomson 1952 / Ex. Musaeo James Thomson  ; Crimea (1♂): cinerarium F.  , Krimm, Rib. / Museum Paris Coll. M. Pic  ; Crimea (1♂): cinerarium Crimée / Museum Paris Coll. M. Pic [ MNHNAbout MNHN]; Tatarbunar  , 3–5.V.1911, leg. Ivankov A. (1♂) [NMENH].

Dorcadion pusillum v. berladense  . Type material. Lectotype ♂ (present designation): 78 / pusillum var berladense Pic  (handwritten label) / Type [red label] / Museum Paris Coll. M. Pic ( Fig. 1AView FIGURE 1) [ MNHNAbout MNHN]. Non-type material. Moldavie, Vall. du Berlad, A. L. Montandon / reçu de a. nicolas / type [handwritten label] / Museum Paris Coll. M. Pic (1♀, Fig. 1BView FIGURE 1); Moldavie, Vall. du Berlad, A. L. Montandon / decipiens  ? ♂ / Museum Paris 1911, Coll. J. Bourgeois / Dorcadion pusillum Küst. Breuning  dét. (1♂, Fig. 1DView FIGURE 1); Moldavie, Vall. du Berlad, A. L. Montandon / pusillum Kust.  / pusillum  d’après Daniel / Dorcadion pusillum berladense Pic, Breuning  dét. / Museum Paris 1911, Coll. J. Bourgeois (1♂, Fig. 1CView FIGURE 1) [ MNHNAbout MNHN]; Moldavie, Vall. du Berlad, A. L. Montandon / Dorc. pusillum Küst., M. Al. Ieniștea  det. (1♂); Moldavie, Vall. du Berlad, A. L. Montandon / Mus. Bucuresci dăruit [gift] A. L. Montandon / Dorcadion decipiens Germ.  , dét Nicolas / Dorc. pusillum Küst. M. Al. Ieniștea  det. (1♂, Fig. 1FView FIGURE 1); Moldavie, Vall. du Berlad, A. L. Montandon / Dorcadion decipiens Germ.  / Dorc. pusillum Küst., M. Al. Ieniștea  det. (1♂) [ MGABAbout MGAB, Nr. 85845, Heritage coll.]; Moldavie, Vall. du Berlad, A. L. Montandon (1♀) [ MGABAbout MGAB, Nr. 199, Serafim coll.]; Moldavie, Vall. du Berlad, A. L. Montandon (2♂♂) [ MGABAbout MGAB, Nr. 206, Serafim coll.].


Museum National d'Histoire Naturelle


Muzeul de Istorie Naturala "Grigore Antipa"


Desert Experiment Station of the W.I.R.














Dorcadion pusillum pusillum Küster, 1847

Dascălu, Maria-Magdalena 2018


Dorcadion pusillum Küster, 1847 : 90

Küster, 1847 : 90


D. pusillum v. berladense

Pic, 1903 : 7


D. pusillum

Breuning, 1946 : 100


Dorcadion pusillum

Kraatz, 1873 : 75 Ganglbauer, 1884 : 450 Montandon, 1908 : 100 Plavilstshikov, 1958 : 149 Panin & Săvulescu, 1961 : 389 Breuning, 1962 : 112 Kasatkin, 2006 : 91 Lazarev, 2009 : 198 Danilevsky, 2010 : 251 Lazarev (2011) : 257


Dorcadion pusillum pusillum

Kasatkin, 2002 : 277 Danilevsky, 2010 : 251


D. pusillum v. berladense

Montandon, 1908 : 100


D. pusillum

Plavilstshikov, 1958 : 149 Panin & Săvulescu, 1961 : 389


D. pusillum

Plavilstshikov, 1958 : 149


D. pusillum

Panin & Săvulescu, 1961 : 389


D. pusillum

Breuning, 1962 : 112


D. pusillum

Breuning, 1962 : 159


D. pusillum berladense

Kasatkin, 2002 : 277 Danilevsky, 2010 : 251