Vipera ammodytes, (Linnaeus, 1758) (Linnaeus, 1758)

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review

The description year of Coluber ammodytes has been arbitrarily fixed as 1758 since the species was indexed in the 10 th Ed. of ‘Systema Naturae’ ( Linnaeus 1758), the starting point of zoological nomenclature ( ICZN 1999: Art. 3). The main description of the taxon was published ten years earlier in a pre-‘Systema Naturae’ work ( Linnaeus & Sundius 1748).

The description on page 216–217 of ‘Systema Naturae’ edition 10 ( Linnaeus 1758) ( Fig. 1 View FIGURE 1 ) translates as follows:

Species numbered 174. [ Coluber ] Ammodytes . 142 ventral scales— 32 subcaudal scales Amoen. acad. I. p. 506 n. 25. (reference to Linnaeus 1749a, P1)

Venomous snake (Mars sign ♂, see Krecsák & Wahlgren 2008). Bellon. itin., 203. Druinus. (reference to Bellon 1605, P2)

Aldr. Serp. 169. Ammodytes. (reference to Aldrovandi 1640, P3)

Lives in the Orient.

Nose terminated in a raised wart.

In the 12 th edition of ‘Systema Naturae’ ( Linné 1766) Linnaeus included the same text ( Fig. 2 View FIGURE 2 ).

Amoen. acad. I p. 506 n. 25 is a reference to ‘Surinamensia Grilliana’ ( Linnaeus 1749a), which was initially published in 1748 as a dissertation ( Linnaeus & Sundius 1748), defended by Peter Sundius (1725–1786) on the 18 th of June 1748. Once a dissertation was published in the compilation ‘Amoenitates Academicae’, which often included alterations from the original text, Linnaeus started referring to this version and dropped the citation of the original dissertation (Krecsák & Bauer in prep.). In this instance, however, the Coluber ammodytes text from the dissertation by Linnaeus & Sundius (1748) was not altered before its publication in ‘Amoenitates Academicae’ ( Linnaeus 1749a). ‘Amoenitates Academicae’ was printed in multiple editions, often with altered text, and selected dissertations were additionally translated in various languages. As such three Latin editions of ‘Surinamensia Grilliana’ can be identified: Ed. 1 printed in Leiden (Lugduni Batavorum) with the dissertation on pages 489–519 ( Linnaeus 1749b), Ed. 2 Stockholm and Leipzig (Holmiae et Lipsie) with the dissertation on pp. 483–508 ( Linnaeus 1749a) and the third Erlangen (Erlangae) edition which included the dissertation similarly on pp. 483–508 ( Linnaeus 1787). The content of the three editions of ‘Surinamensia Grilliana’ is similar. Linnaeus always referred to the Stockholm and Leipzig edition ( Linnaeus 1749a).

‘Surinamensia Grilliana’ included animals that mainly originated from Dutch Guyana (now Suriname) in South America. The animals were collected by the plantation owner Pater Gerret and his son from Surinam and given to Claes Grill (1705–1767) who later donated the collection to the Museum Upsaliense. Grill was a Swedish merchant, factory and ship owner, director of the Grill Trading House, one of the leading companies in the East India trade, who had a particular interest in natural sciences and financially supported Linnaeus. The collection included 24 animals of which 17, numbered 3–19, were Amphibia.

In an appendix to the collection, one snake, COLUBER scutis abdominalibus CXLII, Squamis caudalibus XXXII No. 25, is named Ammodita and described: ‘Illustrandae historiae naturalis gratia, serpentum maxime, illius vero imprimis speciei, quae ab Ammodita nomen tulit, Colubrum subjugam ab Illustri & Nobilliss. Regiae Cancellariae Consiliario, D:no EDVARDO CARLESONIO, cum ad Portam Ottomanicam munus legati Extraordinaii obiret, captum.’ (“To illustrate the grace of natural history, particularly of snakes, and of the particular species, which took its name from Ammodita, a Coluber was subdued by EDVARDO CARLESONIUS the illustrious and most noble counsellor of the royal chancellery, collected, when he was performing the function of extraordinary ambassador at the Ottoman Gate”).

Linnaeus provided two secondary references for the species: Ammodytes ‘Raj.quadrup. 287.’ (S1) and ‘Jonst. Serp. 14.t.I.f.3.’ (S2). He describes the colour pattern of the snake, its scales, mentions that it has two large fangs, and provides its distribution as southern parts of Europe, Italy, Illyria and that it is also found in Libya. He further refers to the horned Aspis (Aspidem cornutum) described by Matthioli (1565) (S3) as found in Italy.

Linnaeus listed Libya as an area of occurrence for V. ammodytes , based on Aldrovandi (1640) and Jonstonus (1657). These authors derived this information from Solinus (1572), who equated Africa to Libya: ‘Porro quod in illo ambitu Aegyptium finitur pelagus et Libycum incipit, placuit ut Africam Libyam diceremus.’ (‘Hereafter, because the Egyptian Sea finishes in this circuit and the Libyan begins, it seems good to me to call Africa Libya.’). However, Solinus only mentioned ‘Ammoditae’ as present in Africa. This name is similar to Hammodytes used by Marcus Annaeus Lucanus ( Anonymous 1592), which rather refers to the Saharan sand viper ( Cerastes vipera ), and it is likely that both authors, who were referring to Libyan snakes, intended the same species.

The specimen collected by Carleson was depicted on the plate that accompanied the dissertation ( Fig. 3 View FIGURE 3 ). The illustrator was P.A. Petersson, and the engraver was Carl Bergquist (1711–1781) who was employed by the Swedish Academy.

Linnaeus published an anonymous review in Swedish of the dissertation in ‘Lärda tidningar’ (Scholarly Newspapers) ( Anonymous 1748). This was not a unique case, multiple instances are documented in which he wrote extensive and highly positive reviews, advertising his publications ( Lindell 2012; Wahlgren 2012; Krecsák & Bauer in prep.).

For a review of the specimens associated with the primary and secondary references, see Table 2 View TABLE 2 .

Although it can be established with certainty that the specimen stored in UPSZMC is the one depicted on plate I of ‘Surinamensia Grilliana’ ( Linnaeus & Sundius 1748; Linnaeus 1749a), we have scrutinized and reviewed the manuscript and published catalogues of the collections in order to identify any further V. ammodytes specimens that might have been studied by Linnaeus.

The type material

The illustrated specimen can be traced throughout the manuscript catalogues of the collection (see Krecsák & Wahlgren 2008). UPSZMC housed a specimen (or several specimens) of Coluber ammodytes , as the species was recorded in Catalogue I, the oldest account of the collection from the 1780s. Catalogues II to VIII recorded one specimen of Coluber ammodytes from the AL (i.e., Alströmer/Linné) donation, and Catalogue IV contains a remark that the specimen was in good condition. Ossian Olofsson’s ‘Linne´samlingarna’ contains the same info as above.

Thunberg (1787) listed the specimen in the ‘6. et 7. Donatio 1749 et sequentibus annis Jonae Alströmer nec non Caroli a Linné’ donation and states that it was donated by Edvardo Carleson, counselor of the king’s chancellery and refers to Linnaeus’s dissertation in ‘Amoenitates Academicae’, giving two scale counts 142:32, and a short comment in a footnote: “Dono datus ab Edvardo Carleson, Reg. Cancellariae Consiliario. LINN. Amoen.Acad. Tom. I p. 517.”. The reference was considered a misprint for page 506 by Lönnberg (1896), but it was actually only a difference in the editions used. Linnaeus used and referred to the Stockholm edition ( Linnaeus 1749a) of ‘Amoenitates Academicae’ where the description begins on page 506, whereas Thunberg used the Leiden edition ( Linnaeus,1749b) in which the Appendix starts on page 517. The scale counts given by Thunberg agree with the counts in different works by Linnaeus (1749a, 1758, 1766). Lönnberg (1896) mentioned that the individual is the same as depicted on plate XVII of ‘Surinamensia Grilliana’ ( Linnaeus 1749a), with the mouth still opened. Wallin (2001) listed the specimen as donated to the museum by C. von Linné, with a note that it originated from E. Carleson.

At present there are no specimens of Coluber ammodytes housed in The Swedish Museum of Natural History that are possible candidates for being material examined by Linnaeus. The collection housed a snake (NRM 784) from the De Geer collection, which was identified by Charles De Geer as Coluber ammodytes . Records show that the particular specimen could not have been present in De Geer’s collection until after 1758. It was reidentified as Xenodon severus by Lars Gabriel Andersson around 1898. The specimen could not be found in the NRM collection in 2024, but there is no indication that it was examined by Linnaeus, and it is not considered as ever being part of the type series.

Primary references P2 and P3 and secondary references S1–S11 contain descriptions of V. ammodytes and/or other snake species, as shown is Table 2 View TABLE 2 . There are no known extant specimens that may have been the source of the above descriptions.

Description of the lectotype

Specimen: UPSZTY 95, Nr. 95, Donation: C. Grill. ( Fig. 4 View FIGURE 4 ).

Printed label by Dr. Åke Holm (1909–1989), former Director of the Zoological Museum, in the jar: Uppsala Univ. Zool. Mus. Linnésamlingen nr. 95 Coluber ammodytes . Information on the card file in the Linnaean collection: Coluber Ammodytes L. ; Linnésaml. 95; Vipera ammodytes Latreille ; Ldb 1897no. 25; Mus. Grill 25. Additionally, the part dealing with the taxon has been cut and glued from a reprint of Lönnberg’s 1896 paper.

The individual is preserved in a new glass cylinder to which it was transferred during the 1950s by Holm (L. Wallin pers. comm.). The original jar with the rectangular paper label with ink inscription in Thunberg’s handwriting ( Fig. 5 View FIGURE 5 ) is stored on a separate shelf with other glassware.

The specimen, a subadult male, shows signs of damage ( Fig. 4 View FIGURE 4 ). Its mouth was probably kept open with a solid object in order to show its fangs. Most probably it was killed with a stick, but the damage was not depicted on the plate. Artistic freedom and the primary aim of depicting the species, not the specimen, motivated the illustrator to draw an intact snake that shows no signs of damage. Examples of similar approaches of artistic freedom while depicting preserved material are well known from the 16 th to 18th centuries ( Simmons & Snider 2012).

Morphometric characters: SVL: 216, TL: 21, L cor: 202.42, Lt cor: 8.82, Alt cor: 8.69, L cap: 13.58, Lt cap: 12.83, Do: 1.91/1.95, Dols: 1.92/1.94, Alt corni: 2.07, Alt r: 2.88, L scr: 2.81, Lt scr: 2.38.

Scalation characters: PreV: 5, Ventr: 149, DorsN: 21, DorsMb: 21, DorsT: 17, Scd: 33/33, SupL: 9/9, SubL: 10/10, Horn: 17, Cant: 2/2, Apic: 2/2, Lor: 6/6, CircO: 10/10, CircO2: 3/5, CircO2C: 2/1, Im/SL:4/4, Gul: 4/4,

Colour pattern and qualitative characters: Very pale currently. Linnaeus ( Linnaeus & Sundius 1748; Linnaeus 1749a) described the specimen as grey and almost orange in colour, with darker dorsal zigzags, ZZW: 36/34, ZZW2: 10/10. Belly and subcaudal surface light beige. States of qualitative traits according to Tomović (2006): I:2, II:0, III:0, IV:1, V:1, VI:3, VII:2, VIII:0, IX:0.

Nomenclatural aspects

Mertens & Müller (1928) proposed the restriction of the type locality of Vipera ammodytes ammodytes (Linné) to Illyria.

Schwarz (1936), in the synonymy of Vipera ammodytes ammodytes , mentioned the name Ammodytes from the work ‘ Serpentum et Draconum Historiae’ by Aldrovandi (1640) and gave the provenance of the individual as Dalmatia , Goerz , Japidiç Mountains . For Coluber ammodytes Linnaeus, 1758 , Schwarz stated that the individual described in the 10 th edition of ‘ Systema Naturae’ ( Linnaeus, 1758) was the specimen previously described by Aldrovandi. He also proposed the restriction of the type locality to Zara (Zadar; Croatia) without clarification or explanation. Later, the restricted type locality, Zadar, was followed by Mertens & Müller (1940), Mertens & Wermuth (1960), Saint Girons (1978), McDiarmid et al. (1999), and Heckes et al. (2005).

Bruno (1968) restricted the type locality to near Castello Nuovo di Duino (Trieste, Venezia Giulia NW), NE Italy, noting the holotype as male uv., 95 SCL-ZMUU, leg. E. Carlesonio ( Bruno 1968 p. 293, plate 1). He discussed the source of the individual, arguing that it was given to Linnaeus by the councillor E. Carlesonio, ambassador of the Republic of Venezia in Turkey (Türkiye), who found it on a pile of sand near the entrance to the Duino Castle. This specific demarcation of the collection place could not be found in any printed material, and the source could not be obtained from the author (Bruno pers. comm.). Bruno performed an assessment of the primary references in ‘Systema Naturae’ edition 10 ( Linnaeus 1758), reviewed the origin of the species and specimens described therein and thereafter unambiguously referred to the specimen UPSZTY 95 as the typus, thereby designating the specimen as the lectotype in accordance with Article 74.5 of the Code ( ICZN 1999).

Heckes et al. (2005) questioned the validity of Bruno’s (1968) decision and argued that the only certain information is that provided in the original description ( Linnaeus 1758), i.e. ‘Habitat in Oriente’, and considered the restriction of the type locality to Zadar, proposed by Schwarz (1936) to be correct, although the logic of this is unclear.

Results of the statistical analysis

Morphometric and meristic characters: Results of Discriminant Canonical Analysis on 12 morphometric and meristic characters showed that, in the projection of the first and second canonical axes, the lectotype of Vipera ammodytes falls within the 95% confidence intervals of the multivariate variability of two subspecies ( V. a. ammodytes and V. a. montandoni) ( Fig. 6 View FIGURE 6 ).

Qualitative characters: Frequencies of qualitative traits of the subspecies and lectotype are given in Table 3 View TABLE 3 .

Correspondence analysis of four taxonomically informative qualitative characters (sensu Tomović 2006) showed that in the projection of the first and the second correspondence axes the lectotype specimen fits with the specimens of the V. a. montandoni subspecies ( Fig. 7 View FIGURE 7 ).

These are characterized by the following characters and combination of states: IV:1 (greater height of rostral plate) and V:1 (greater height of rostral plate than of nasorostral plates). This subspecies is also characterized by the more frequent presence of the character state I:0 (suprarostral plate lacking), while the lectotype has I:2 (two suprarostral plates) and II:0 (neither nasorostral plate in contact with canthus rostralis), which are also frequent in both V. a. montandoni and V. a. meridionalis (see Table 3 View TABLE 3 ).

Vipera a. meridionalis is additionally characterized by the more frequent presence of character state I:3 (three suprarostral plates) and V:0 (heights of rostral and nasorostral plates equal) ( Table 3 View TABLE 3 ).

In contrast, specimens from the range of the nominotypic subspecies (V. a. ammodytes ) are characterized by the more frequent presence of the following character states: I:1 (one suprarostral plate), II:1 (both nasorostral plates in contact with canthus rostralis), IV:2 (greater width than height of rostral plate) and V:2 (greater height of nasorostral plates than of rostral plate) ( Table 3 View TABLE 3 ).