Coluber berus

Krecsák, László & Wahlgren, Richard, 2008, A survey of the Linnaean type material of Coluber berus, Coluber chersea and Coluber prester (Serpentes, Viperidae), Journal of Natural History 42 (35 - 36), pp. 2343-2377 : 2352-2358

Coluber berus

We could not identify any specimen labelled berus as a species designation with possible type status in the Stockholm collection, i.e. NRM. However, UUZM houses a single specimen with possible type status labelled C. berus ( Figure 1 ).

Description of the specimen

Specimen. UUZM 1 , Donation: C. Gyllenborg. Printed label by Holm on the actual jar: Uppsala Univ. Linnésamling Nr 1 Mus. Gyll. nr 1. Printed label by Holm in the jar: Uppsala Univ. Zool. Mus. Linnésamlingen nr. 1 Coluber berus . Original jar with Thunberg’s label is lost. Female .

Mensural characters. SVL: 234, TL: 22, HL: 11.86, HW: 8.92, HD: 4.96, DbE: 5.86, RML: 14.37, DbER: 4.43/4.37, DbES: 1.47/1.46, EyeHD: 2.20/2.29, EyeVD: 1.94/ 1.83, FL: 2.18, FW: 1.95, RL: 2.02, RW1: 0.97, RW2: 2.12, Nas: 1.89/1.90, Nos: 0.87/0.97, DbNasSupL: 2.19/2.17, DbNosSupL: 1.35/1.26, DL: 2.20 (all measurements are in mm).

Scalation characters. PreV: 5, Ventr: 144, DorsN: 21, DorsMb: 21, DorsT: 17, DR23: 7, DR21: 9, DR19: 69, DR17: 104, Scd: 31/31, SupL: 8/8, SubL: 10/10, Lor: 2/3, CircO: 9/8, SubLSubO: 4–5/4–5, Apic: 2, Canth: 2/2, IntC: 7, IntSupO: 3/2, SbE: 12, Crws: 17, GulR:3/4, GulT 20/22. Undivided rostral scale; nasal scales separated from the rostral by one nasorostral scale; nasals not connected to the circumocular scales; undivided nasals; nostrils in the upper part of the nasal scales; circumocular scales in direct contact with the supralabial scales; undivided supraoculars; subcaudal scales divided.

Colour pattern. Very pale; most probably entirely dark belly; head pattern once formed by two separated dark oblique bands forming a barely visible inverted V mark on the head; dorsal pattern formed by an undivided, sharp tipped, zigzag band, consisting of 82/– ZZW on the body and 17/– ZZW2 on the tail.

Review of the literature

Linnaeus (1758: 217) described the species in Systema Naturae as Coluber Berus , species number 183. The number set ‘146–39’ represents the key characters for the species, i.e. the number of ventral and subcaudal scales. Three sources are cited: Faun. svec. 260 ( Linnaeus 1746b), Amoen. acad. I. p. 113. n. I ( Linnaeus 1749a), and Aldr. serp. 115, 116 ( Aldrovandi 1640). Linnaeus employed the Mars sign „ to denote venomous snakes (now used to designate the male sex) in several works (e.g. Linnaeus 1754, 1758, 1761, 1766) but explained its use only in Systema Naturae ( Linnaeus 1758: 221, 1766: 382) ‘‘ Telo venenato reflexili armatos, signo „ notatos, observa. Translation by Kitchell and Dundee (1994): Note those armed with a retractable, venomous weapon, marked by the symbol „.’’ Linnaeus indexed this specimen in two subsequent works: Fauna Svecica ( Linnaeus 1761: 104), species 286, and Systema Naturae ( Linnaeus 1766: 377), species 183.

The same specimen was previously described in detail by him in Amphibia Gyllenborgiana ( Linnaeus 1745a: 5–6, 1749a: 113–114), as specimen a of ‘I. Coluber Scutis abdonminalibus CXLIV, caudalibus XXXIX ’. From the relatively high number of subcaudal scales we can assume this is a male. Presumably it is the specimen mentioned in his later works as C. berus (1746b, 1758, 1761, 1766). We can suppose that Linnaeus made a copy error when quoting the number of ventral scales from this work, or from his copy of Fauna Svecica (1746b), to Systema Naturae (1758). When compiling the subsequent works (1761, 1766) he used the description in Systema Naturae (1758) and not the description in earlier works. Linnaeus described the scales on the head and body of the specimen, its fangs and remarked that it is completely black.

Linnaeus (1745a: 6; 1749a: 114) also described another specimen, B, as ‘‘I. Coluber Scutis abdonminalibus CXLIV, caudalibus XXXIX ’’. This was a normal coloured male specimen, possessing a dark zigzag band. Linnaeus also described its hemipenis, but interestingly gave no scale numbers, only mentioned that the snake has numerous ventral and subcaudal scales. In each case where Linnaeus described a specimen he had studied, he recorded these two scale numbers, thus this description may be based on an artistic source. From the four references ( Linnaeus 1742; Petiver 1695; Ray 1693; Seba 1735), the latter, cited as ‘ Vipera vera indiae orientalis. Seb. thesaur. t. II. p. 9. tab. 8’ is of significance. Seba (1735: 9, pl. 8, f. 4) depicted a male adder-like specimen with erected hemipenis (specially marked with the letter A) and described it as ‘ Vipera , vera, Indiae Orientalis ’. Though we are unable to positively identify the snake depicted on the Seba plate, it is most probably a V. berus .

Linnaeus (1746b: 97 no. 260) described in Fauna Svecica the Coluber a from Amphibia Gyllenborgiana ( Linnaeus 1745a, 1749a). The number of ventral and subcaudal scales given was 144 and 39 with the citations Amph. gyllenb., implying this dissertation.

In the sixth edition of Systema Naturae Linnaeus (1748: 34) listed another specimen, which identification he based on the scale counts. He recorded it with No. 20 under Coluber as ‘Fn, 260 Vipera’ (Linnaeus 1746: 97) and possessing 145 ventral and 36 subcaudal scales. Linnaeus (1749c) later cited this reference in the main description of Coluber chersea in order to corroborate the difference in the number of scales between C. chersea and C. berus . He subsequently referred to the same specimen in the lectures of zoology that were recorded between 1748 and 1752 ( Lönnberg 1913: 157–158).

Male adders are reported in two other pre-Linnaean works, Animalia per Sveciam observata ( Linnaeus 1742) and Öländska och Gothländska Resa ( Linnaeus 1745b: 23), but he provided no details as to whether or not he collected the specimens. The absence of scale counts suggests that he did not.

Works that we should mentioned here, as Linnaeus referred to them in his dissertations and thus the specimen(s) described therein are part of the syntype series, are: Clusii (1605) cited as ‘ Bell. itin. clus. 123. Vipera .’ ( Linnaeus 1746b); Gessner (1621) referred to as ‘ Gesn. serp. 124. Vipera .’ ( Linnaeus 1745a, 1746b); Jonstonus (1657) referred to as ‘ Jonst. serpent. pag 7 ’ ( Linnaeus 1745a); Aldrovandi (1640) referred to as ‘ Aldr. Serp. 115, 116 ’ ( Linnaeus 1758, 1766) and ‘ Aldr. serp. 108. Vipera ’ ( Linnaeus 1745a, 1746b); Charleton (1668) cited as ‘ Charl. onom. 30. Vipera ’ ( Linnaeus 1746b, 1761); Ray (1693) cited as ‘ Raj. quadr. 285. Vipera ’ ( Linnaeus 1745a, 1746b, 1749a 1761); Petiver (1695) cited as ‘ Pet. mus. 17. n. 103. Vipera anglica fusca , dorso linea undulata nigricante conspicua ’ ( Linnaeus 1745a, 1746b, 1749a, 1761); and ‘ Vipera anglica nigricans . Pet. mus. pag. 17. n. 104 ’ ( Linnaeus 1745a, 1749a).

The specimens are the ‘Vipera’, a collective term used for the venomous snakes occurring in Europe, Asia and North Africa in the works by Bellon ( Clusii 1605), Gessner (1621), Jonstonus (1657) and Aldrovandi (1640: 108). Aldrovandi’s plates (115, 116) depict two vipers, which probably are V. aspis , but they are not detailed and we cannot identify them positively. The ‘Vipera’ of Charleton (1668) and Ray (1693) are most probably V. berus . Petiver’s (1695) Vipera anglica fusca , dorso linea undata nigricante conspicua was a normal coloured V. berus inhabiting the woods and dry banks of England, whereas Vipera anglica nigricans was the black coloured morph of the adder. Linnaeus (1761) recognized it later as a separate species, Coluber prester.

Review of the catalogue data

UUZM catalogues. The first inventory of the Gyllenborg donation, which was compiled in 1744, mentioned that there were three ‘viperae’ ( Holm 1957: 25). C. Berus was present in the collection around 1780, as the species is listed in Catalogue I but the catalogue does not indicate the number of specimens.

One specimen in the Gyllenborg donation was recorded in Catalogues II, III, IV and VI. Another juvenile, designated B, and two other specimens in the Thunberg donation appear in Catalogues III, V, VI, VII and VIII. In the latter catalogue one specimen is listed as being black (niger) .

Thunberg (1787b: 8) indexed one Coluber Berus specimen in Donation Caroli Gyllenborg, and two in the Thunberg donation: Coluber Berus : niger 144: 39, and Coluber Berus var. 144: 40 ( Thunberg 1787a: 30). Based on the scale numbers he provided, we deem both were male adders.

Lönnberg (1896: 5) mentioned one specimen in the Gyllenborg donation, with the only remark that it is rather a juvenile form. When Lönnberg studied the material, the label by Thunberg should have been present on the jar, as he stated that the Coluber Berus specimen was labelled ‘ Museum Gyllenborg ’.

Holm (1957: 28) listed the specimen presently numbered UUZM 1 in the Gyllenborg donation and stated that the species is Coluber Berus in the tenth edition of Systema Naturae ( Linnaeus 1758) but that it is not the type. He did not explain the reason for this assumption. On the card file of the Linnaean collection Holm provided the following additional information to the specimen ‘‘ Is probably the specimen, which in Thunberg’s catalogue is designated as ‘Col. Berus B pullus Th’ and is probably not part of the collection ’’. Wallin (2001: 123) mentioned the existence of the specimen, and recorded it as not being a type referring to Holm (1957).

NRM catalogues. Coluber berus specimens were present in the Royal Academy’s collection, as the taxon was recorded in Catalogue IX by Hornstedt. The pencil note ‘v 3 ex. ’ by Quensel, probably indicates that three specimens were in the collection .

‘ Catalogus Amphibiorum …’ (Catalogue X) by Quensel contains two C. berus specimens, one with right colouration and another, with a mixture of C. prester and C. chersea colour pattern (probably dark brown).

Catalogue XI, compiled by Quensel listing the duplicates, contains one C. berus specimen that subsequently has been discarded. It could have been sent to the museums in Lund, Uppsala or Åbo ( Finland). The specimen, however, could neither be traced in Lund ( Wahlgren 1865; Rosén 1905; Thomas Madsen, University of Wollongong, personal communication), nor in UUZM. The collection of the Åbo Academy, except a few stuffed birds and mammals, was destroyed in a fire in 1827 ( Fernholm and Wheeler 1983). Similarly, Fernholm and Wheeler (1983) could not trace fishes disposed by Quensel to Uppsala, Lund and Åbo.

Swartz, in Catalogue XII, listed three C. berus specimens, two from the old collection and another juvenile listed at the end of the catalogue and mentioned the acquisitions in 1810. Dalman in both Catalogues (XIII and XIV) mentioned the taxon in the collection, but they do not contain any information on the number of specimens present. Catalogue XIV reports Vipera Berus and Vipera Berus obscurior in the collection. The latter name could have been used for specimen(s) Dalman was unable to assign to any of the three Coluber .

Andersson (1899) did not mention C. berus among the Linnaean specimens.

Discussion

Based on the literature surveyed we presume that Linnaeus examined only two specimens, both V. berus : a primary syntype, the male a, possessing 144 ventral and 39 subcaudal scales, in Amphibia Gyllenborgiana ( Linnaeus 1745a, 1749a) and later indexed in several works by Linnaeus (1746b, 1758, 1761, 1766), and a further male, possessing 145 ventral and 36 subcaudal scales, indexed in Systema Naturae ( Linnaeus 1748) and mentioned later in his lectures ( Lönnberg 1913).

The UUZM catalogues prove the existence of a specimen in the Gyllenborg donation. Three other C. Berus specimens were housed during the 18 th century in the collection, two donated prior to 1787, as they were mentioned by Thunberg (1787a), and another donated later.

UUZM, at present, houses five V. berus specimens, which were once part of the Museum Thunberg. One melanistic and a normal coloured V. berus with dark ground colour are catalogued as 56254 (old no. 435) Vipera berus var. Museum Thunberg. The melanistic juvenile male possesses 142 ventral scales and 40/40 subcaudal scales and it might fit Thunberg’s Coluber Berus : niger. This should have been the specimen B in the manuscript catalogues. The other specimen, under UUZM 56265, is a normal coloured male that has 144 ventral scales and 38/38 subcaudal scales. This could be the other specimen listed by Thunberg in his printed catalogue ( Thunberg 1787a: 30), and numbered as c or d in the manuscript catalogues. The third specimen in the manuscript catalogues might be one of the three V. berus specimens labelled as Museum Thunberg in UUZM under the catalogue no. 56262 (old no. 442): one normal coloured juvenile female, with 150 ventrals and 33/33 subcaudals; a normal coloured juvenile male with 150 ventrals and 44/43 subcaudals, and another normal coloured juvenile male possessing 147 ventrals and 38/40 subcaudals. Nothing indicates that any of these specimens could be primary, secondary or tertiary syntypes.

The specimen indexed UUZM 1 is not a melanistic specimen, thus does not agree with specimen a presented in Amphibia Gyllenborgiana ( Linnaeus 1749a). Additionally, it is a female possessing only 32 subcaudal scales. Linnaeus recorded 39 subcaudal scales on the male specimen he described and we can assume that he would not by mistake have counted this significantly lower number of scales; therefore we can not assess it as a syntype.

Six old adder specimens have been identified in NRM that might have been listed in the catalogues above. Three specimens were probably collected or donated during Dalman’s time, as they bear labels ( Vipera Berus ) by Dalman only: specimens NRM 195 and NRM 6093 with unknown origin and one specimen, NRM 6095, collected by Dalman. A dark-coloured specimen NRM 30204, labelled by Dalman ‘ Vipera Beru s. Var. Obscura . Dalm.’ Originates from the collection of Gustaf von Paykull, and was probably listed as Vipera Berus obscurior in Catalogue XIV. Two specimens are catalogued as NRM 6094 (old numbers 550008 and 550009). The jar bears a label, ‘ Ber.’ by Quensel and a label by Dalman ‘ Vipera Berus et Chersea ’. The specimens were probably mixed by Dalman. In the opinion of Andersson (written on his label), these are the two Vipera Berus in Catalogue X by Quensel. Nothing indicates that any of these specimens was ever studied by Linnaeus and could have syntype status.

Taxonomic and nomenclatural aspects

Mertens and Müller (1928) proposed the restriction of the type locality of V. b. berus to Sweden. Schwarz (1936) proposed the change of the type locality from Sweden to Uppsala, a restriction proposed once again by Mertens and Müller (1940), apparently by oversight. The restriction of the type locality to Uppsala has been attributed to Mertens and Müller (1940), not to Schwarz by several authors: Mertens and Wermuth (1960), Saint Girons (1978), Bruno (1985), Golay et al. (1993), Nilson et al. (2005). These restrictions were not based on the original type series, as the authors merely proposed them without further comments. According to Art. 76.1 and Art. 73.2.3 ( ICZN 1999) they should not be considered valid. Golay et al. (1993) stated that the type of V. berus is lost, whereas according to McDiarmid et al. (1999) the specimen UUZM Nr. 1 preserved in UUZM is the holotype, but we have shown that this is not Linnaeus’s original specimen and therefore this conclusion is erroneous.

As Linnaeus referred to several literature sources and described further specimens, we have a syntype series which consisted of one primary syntype, a black male ( Linnaeus 1745a, 1746b, 1749a, 1758), two untraced, male, secondary syntypes recorded by Linnaeus (1742, 1745a, 1746b, 1749a), specimens described and depicted by Aldrovandi (1640), and several tertiary untraced syntypes described and depicted by Clusii (1605), Gessner (1621), Jonstonus (1657), Charleton (1668), Ray (1693), Petiver (1695) and Seba (1735). The vipers in some of the literature sources cited are collective names or species other than V. berus . This does not cause nomenclatural instability, as all specimens described by Linnaeus were certainly V. berus and this taxon was well defined by the author.

Based on the available information the type status of the specimen UUZM 1 cannot be assessed, but it is not the primary syntype .

For reason of stability we consider it appropriate to designate a neotype and restrict the type locality according Art. 76.2 ( ICZN 1999 ) to Berthåga , Uppsala (59 ° 519N 17 ° 349E), Uppland , Sweden. This is a well known locality for adders, in the northern part of Uppsala, that even might have been visited by Linnaeus, and where the species still occurs (Mats Eriksson, UUMZ, personal communication) .

Description of the neotype

Specimen. UUZM, Nr. 7454; Berthåga , Uppsala (59 ° 519N 17 ° 349E), Uppland, Sweden; 02.06.1934; coll. Anonymous. Adult male (erected hemipenis) ( Figure 2 ) .

Mensural characters. SVL: 390, TL: 65, HL: 18.83, HW: 12.50, HD: 7.47, DbE: 8.17, RML: 22.99, DbER: 5.73/5.79, DbES: 4.92/5.00, EyeHD: 3.09/3.07, EyeVD: 2.65/ 2.61, FL: 4.19, FW: 2.66, RL: 2.81, RW1: 1.34, RW2: 3.62, Nas: 2.76/2.84, Nos: 1.21/1.18, DbNasSupL: 3.29/3.35, DbNosSupL: 1.78/1.82, DL: 3.58 (all measurements are in mm).

Scalation characters. PreV: 4, Ventr: 138, DorsN: 23, DorsMb: 21, DorsT: 17, DR23: 11, DR21: 11, DR19: 105, DR17: 121, Scd: 41/41, SupL: 8/8, SubL: 11/11, Lor: 2/3, CircO: 11/10, SubLSubO: 4–5/4–5, Apic: 2, Canth: 2/2, IntC: 9, IntSupO: 3/2, SbE: 10, Crws: 18, GulR:5/5, GulT 31/34. Undivided rostral scale; nasal scales separated from the rostral by one nasorostral scale; nasals not connected to the circumocular scales; undivided nasals; nostrils in the middle of the nasal scales; circumocular scales in direct contact with the supralabial scales; undivided supraoculars; subcaudal scales divided.

Colour pattern. Dark gray background colour; entirely dark belly; head pattern formed by two separated dark oblique bands forming an inverted V mark on the head; dorsal pattern formed by an undivided, sharp tipped, dark zigzag band, consisting of 75/76 ZZW on the body and 25/21 ZZW2 on the tail.

Figures

Figure 1. Specimen UUZM 1, labelled Coluber berus.

Figure 2. The designated neotype of Coluber berus, specimen UUZM 7454.