Scolopendra oraniensis Lucas, 1846

Doménech, Carles, 2024, Taxonomic and nomenclatural reassessment of the Iberian Peninsula’s nomina obscura, Scolopendra viridipes Dufour, 1820 and S. chlorotes L. Koch in Rosenhauer, 1856 (Chilopoda, Scolopendromorpha, Scolopendridae), ZooKeys 1208, pp. 49-80 : 49-80

Scolopendra oraniensis Lucas, 1846

Types: Figs 7 , 8 ; Table 3 . Non-types: Figs 1 C – I , 2 B, D , 3 , 4 , 5 , 6 , 7 , 8 , 9 ; Tables 1 , 2 , 3 ; Suppl. material 1: files 1, 3

Scolopendra viridipes Dufour, 1820 ; nomen oblitum; senior syn. nov.

S. oraniensis Lucas, 1846 ; nomen protectum.

S. chlorotes L. Koch in Rosenhauer, 1856; junior syn. nov.

S. mediterranea Verhoeff, 1893: 318 .

S. mediterranea lusitanica Verhoeff, 1893: 318 .

S. clavipes Silvestri, 1897: 7 (sic.).

S. oraniensis africana Attems, 1902: 556 .

Rhadinoscytalis canidens oraniensis : Attems 1926: 246.

S. canidens oraniensis : Attems 1930: 19, 36, 37, fig. 50.

S. canidens lusitanica Verhoeff, 1931: 309.

S. oraniensis : Würmli 1980: 348–350.

Morphological diagnosis

[based on S. oraniensis type series]. Body length up to 64 mm. 18 or 19 antennal articles; 5 ½ basal ones glabrous in their entire surface. Antennae / cephalic plate length ratio ≈ 3.30. Forcipular trochanteroprefemoral process clearly defined, with two or three rather inconspicuous denticles. Tooth plate with 4 + 4 (rarely 3 + 3) teeth, divided into two groups. T 1 without sutures or sulci. Paramedian sutures complete on tergites TT 2–20. T 21 with a complete median longitudinal suture. Margination starting at T 14, complete on TT 19–21. Sternite paramedian sutures on TT 2–20. Coxopleuron basally pore-field, with a single medio-distal spine. Coxopleural process with a small pore-field, sub-cylindrical and quite elongated, with 9–14 spines altogether, disposed in sub / apical, dorsal, or lateral positions. Legs 3–20 with few setae. Two tarsal spurs on legs 1; a single tarsal spur on legs 2–18 or 19. Ultimate leg elongated, sometimes with a sinusoid transverse sulcus on the ventro-distal part; prefemoral spines (usually between 18–28) arranged in five frequently miss-aligned or duplicated rows with the VL: 1-5, V: 2-6, VM: 2-6, M: 4-8 and DM: 2-4 formula. Prefemoral process inconspicuous ending with two or three (rarely four) spines. Prefemur and femur of UL glabrous; tibia distally covered by sparse setae; tarsi 1 and 2 covered by small setae. UL / T 21 length ratio ≈ 5.15.

Etymology.

from the toponym “ Oran ”, Algeria and the feminine (or masculine) suffix - ensis (from) meaning “ from Oran ”, Algeria, in reference to the type locality.

Type series and type depository.

Lucas (1846) did not declare on which specimens he based the nominal taxon of S. oraniensis (type series) ( ICZN 1999: Art. 72.4. 1), neither exposing the specimens’ depository nor designating a holotype ( ICZN 1999: Art. 72.1. 1). Otherwise, Kraepelin (1904) indicated the presence of the types [syntypes] at the MNHN in writing “ – [ Scolopendra ] oraniensis Lucas. – Algérie: envir. d’Oran [surroundings of Oran] (H. Lucas, 1849 [sic.]. Types) ”, without further data. Therefore, after the examination of the MNHN specimens and their labels (Figs 7 , 8 ) ( ICZN 1999: Art. 72.4. 2), I conclude that the type series for the S. oraniensis nominal taxon was erected on the basis of seven syntypes, all of them unsexed adults (Table 3 ). Depository MNHN, Paris, France. Jar Number 282. Samples (Figs 7 , 8 A – C ) and labels (Fig. 8 D ) separated in three different assay tubes (containing 2 / 2 / 3 specimens, respectively).

Collector and collection date.

P. H. Lucas, during winter, between 1839 and 1842 ( Lucas 1846).

Type locality and habitat

from Lucas 1846: Ravines of the “ Sauton’s ” (sic.) [Santon] mountains and ravines between Oran and Mers-el-Kebir, Oran wilayah; Algeria ( ICZN 1999: Recommendation 76 A. 2). Lapidicolous.

Distribution.

Known from southern France (including Corsica), southern Italy (including Sardinia and Sicilia), Malta, Spain (including Balearic Islands), Portugal, Morocco, and Algeria. Introduced in Japan ( Attems 1930; Würmli 1980; Bonato et al. 2017 a).

Proposed new nomenclatural status.

Nomen protectum.

Type series composite redescription

(Table 3 ). Colouration: Colouration of specimens preserved in ethanol is toasted to pale yellow (Figs 7 , 8 A – C ). According to Lucas (1846), colouration of living specimen was as follows [morphological traits are interpreted in brackets]:

(Original colour description available from: https://www.biodiversitylibrary.org/page/2362627#page/301/mode/1up)

Upper body part [anterior tergites] coppery black, lower [posterior tergites] green, in the middle ornamented with a yellowish green longitudinal stripe [probably, the translucence of the Malpighian tubule], [...] jaw [forcipules] reddish [...] palps greenish [second maxillae]; base of antenna green, in the middle greenish and in front stained dark red; feet [locomotory legs] green with dark red nails [unguis proper]; last pair of legs [UL] dark green [...].

Morphological description of the type series.

[notes on brackets are comprehensive annotations from this text author]: Body length up to 64 mm. Antennae reaching up to T 2 [maybe up to T 3; actual length shrivelled by ethanol retraction]; with 18 or 19 articles, the basal 5 ½ ones dorsally and ventrally glabrous (Fig. 7 A, B ). Antennae / cephalic plate length ratio: 2.01 [reduced because of the antennae articles retraction caused by the preservation in ethanol; estimated to be ~ 3.30].

Cephalic plate with disperse puncta and a short anterior longitudinal depression; disto-median or / and paramedian sutures are absent. Posterior edge of cephalic plate overlapping the T 1 (Fig. 7 A ). Coxosternal surface with disperse puncta, without sutures (Fig. 7 B ). Article 2 of second maxillary telopodite with spur. Forcipular trochanteroprefemoral process with one apical and two (rarely one) medial poorly differentiable teeth (Fig. 7 C ). Tooth plates slightly wider than tall, forming an obtuse angle (> 120 °) with respect to the coxosternite; sensillae present. Each tooth plate with 4 + 4, rarely 3 + 3 teeth: the external one separated from the other three and inner ones sometimes fused. (Fig. 7 C ).

Spiracles triangular with three valves, present on body segments 3, 5, 8, 10, 12, 14, 16, 18, and 20.

T 1 without sutures or sulcus (Fig. 7 A ); TT 2–20 with complete paramedian sutures (Fig. 7 D ); T 21 with a longitudinal median suture, surface of all tergites smooth, without depressions (Fig. 8 A ). T 21 width / length ratio ≈ 1.35. Margination starting at T 14, being complete in TT 19–21.

Sternites with complete paramedian sutures from TT 2–20 (Fig. 7 E ). Sternite of ultimate leg-bearing segment (Fig. 8 B ) with sides converging posteriorly; surface smooth, without depressions.

Coxopleuron not surpassing the posterior border of the tergite of the ULBS (Fig. 8 B ); with a dense pore-field area at the base and a single medio-distal spine. Coxopleural process sub-cylindrical and distinctly elongated, reaching up to the first 1 / 4 of the UL prefemoral length. The complete surface is covered by a loose pore-field with some small setae (Fig. 8 C ) and a total of 9–14 usually asymmetrically disposed spines; 4–6 of them in an apical / subapical position; 3–6 in a lateral position, and 1–3 in a dorsal position.

Leg 1 with two distal tarsal spurs on tarsus 1, one lateral anterior and one ventral; legs 1–18 or 19 (mode 19, Table 3 ) with only one ventral tarsal spur. Tibial spurs absent. Legs 1, 2, or 3 with or without isolated setae. Legs 3 or 4–20 with scarce setae.

UL moderately long and slender with ratios of lengths of prefemur and femur = 1.15, femur and tibia = 1.25, tibia and tarsus 2 = 1.80; tarsus 1 and tarsus 2 = 1.20 (Fig. 8 C ). Prefemora flattened dorsally, sometimes with a visible sinusoid transverse sulcus on the ventro-distal side (Fig. 8 A, B ). Spines are variable in number (total: 17–26) and size, usually arranged in five frequently asymmetrical, duplicate, and non-ordered rows, with a formula: VL: 3-6, V: 2-6, VM: 2-6, M: 3-8 and DM: 2-4. VM and M spine rows usually converging proximally; between them a flattened surface is present, regularly rounded proximally and without spines. Prefemoral process inconspicuously prominent, sometimes scarcely setose, ending with two, or rarely three, spines (Figs 8 A – C ). Setae on prefemur and femur almost absent, tibia distally scarce setose and tarsi 1 and 2 densely covered with short setae. UL / T 21 length ratio ≈ 4.39 [reduced because of the retraction of the articles caused by the preservation in ethanol; ~ 5.15].

Genitalia retracted in the whole type series.

Genitalia description.

(Based on S. viridipes and S. chlorotes “ topotypical ” material; Fig. 9 A – D ).

Genitalia well developed, reaching the apical part of coxopleural process when extended. Sternite and tergite of genital segment 1 (TGS and SGS I) convex and distally round, with a proximal median suture – distally attenuated – forming the vertex of a poorly angulated keel (Fig. 9 A, B ). In males (Fig. 9 A ) tergite of genital segment 2 (SGS II) small and horseshoe-shaped, with a poorly visible longitudinal suture. Gonopods short and small. Penis present, with fee endpoint, ventally serrated. TGS, SGS I, SGS II, lamina adanalis, gonopods and penis carrying scatted small setae, the remaining part of the genital structures glabrous. Sexual dimorphism or secondary sexual characteristics indistinct (Figs 9 C, D ); UL tibia and tarsi weakly and inconstantly more hirsute in females, with gender differences frequently indistinguishable.

Species variability.

Scolopendra oraniensis has been observed to be somewhat variable in some few morphological features (Figs 3 – 9 B ; Table 3 , Suppl. material 1: files 1, 3): number of antennal articles (17–20; mode 18–19), number of hirsute basal antennal articles (5–5 ½), number of teeth on tooth plates and on forcipular trochanteroprefemoral processes (respectively 3 + 3 vs 4 + 4; mode 4 + 4 and 3 vs 2; mode 3), beginning of complete margination of tergites (on TT 17, 19, or 20; mode 17), tarsal spurs on leg 1 (0-2; mode 2), single tarsal spurs on legs (2–18 or 19; mode 2–19), coxopleural process morphology and UL prefemur number of spines (respectively 9–14 and 18–32), visibility of UL prefemoral ventro-distal horizontal suture, coxopleural and prefemoral process lengths or number of setae on the UL tarsi.

Additionally, the colouration has proven to be variable sympatrically (Figs 1 D – I , 3 – 9 ). The cephalic plate and tergites can be monochromatic pale yellow to dark green / brown, sometimes reddish, sometimes with a darker / lighter longitudinal stripe. The forcipules can be yellowish to reddish, while the legs and antennae can be pale green, yellowish, bluish, reddish, or bicoloured, with occasional metallic reflections (Figs 1 D – I , 3 , 9 ).

These morphological and colouration characteristics, although tending towards a certain geographic distribution, often turn out to be variable within those populations (Table 3 , Suppl. material 1: files 1, 3). Therefore, the separation of the species by their old names are clearly unjustified, since all the features can be explained by the known intraspecific variability of the taxon ( Iorio and Geoffroy 2006; García-Ruiz 2018), which is also shared with many other species in the genus ( Zalesskaja and Schileyko 1992; Siriwut et al. 2016; Doménech et al. 2018; Dyachkov and Nedoev 2021; Tsukamoto et al. 2021).

Differential diagnosis.

Morphologically, S. oraniensis is related to five other species of the S. canidens group distributed around the Mediterranean Sea and Middle Asia ( Würmli 1980; Simaiakis and Mylonas 2008). Scolopendra clavipes (Middle Asia) can easily be distinguished from S. oraniensis by the clavate morphology of the UL, while the two subspecies of S. dalmatica C. L. Koch, 1847 (northeastern Mediterranean) can be differentiated by their larger size (up to 80 mm), the incomplete median suture of T 21, and the four spines of the UL prefemoral process. However, the morphologically closest relatives to S. oraniensis are S. canidens (southeastern and eastern Mediterranean) and S. cretica Lucas, 1853 (eastern Mediterranean, probably endemic to Crete, Greece). These two latter species can be unequivocally differentiated from S. oraniensis by the number of glabrous articles in the antennae (10–12 vs 5–5 ½, respectively), the type of transition between the glabrous / hirsute articles (gradual vs abrupt), and the number of spines in the prefemoral process (generally 3 vs 2 or 3). Between S. canidens and S. cretica , the only visible difference is the presence of brush-like setae in tarsi 1 and 2 of the UL in the S. cretica females ( Würmli 1980; Lewis 2010; Dyachkov ( ASU) pers. comm. 2022; Huesca pers. comm. 2018).

In these last two species closest to S. oraniensis , comparative analyses such as antennae / cephalic plates and T 21 / UL length ratios and their genitalia descriptions have not been performed yet.

Nomenclatural considerations.

To verify the actual identities of S. viridipes and S. chlorotes , establishing the S. oraniensis specimens Lucas (1846) on which he based the type series was necessary, to have reference material with which to unambiguously compare new material ( ICZN 1999: Preamble, Art. 13, Recomm. 13 A). The S. oraniensis lectotype designation was found to be not nomenclaturally required ( ICZN 1999: 11; point 6 in introduction; Art. 72.1. 1; 74.7). However, neotypes designations for S. viridipes and S. chlorotes were mandatory to clarify their type localities and taxonomic situations in the absence of the original types ( ICZN 1999: Art. 75, 76). Finally, the identities of S. viridipes , S. chlorotes , and S. oraniensis have been solved, and they are all conspecific (Figs 3 – 8 ; Table 3 ) leaving the name S. oraniensis Lucas, 1846 as nomen protectum with which to unambiguously refer to “ all ” of these species ( ICZN 1999: Art. 23.3 and 23.9. 2).

At a nomenclatural level, the specific epithet of S. oraniensis obviously conformed to the reference of the type locality ( Gervais 1847). However, that noun is composed by the feminine (or masculine) suffix “ - ensis ” (coming from) and the prefix (a toponym) “ Orani ”, in the place of “ Oran ”. Coincidentally, “ Orani ” is the name of two other localities located in Italy and the Philippines. This can lead to etymological confusion, especially for the Italian Orani, on Sardinia, a region where S. oraniensis also lives ( Würmli 1980; Bonato et al. 2017 a). Therefore, it is thought to be an incorrect original spelling ( ICZN 1999: Art. 32.4) and according to the author’s intention to honour Oran, Algeria, rather than Orani, Italy, the proper name for S. oraniensis should be amended as “ S. oraniensis ”. However, ICZN (1999: Art. 32.5. 1) clearly states that this supposition should not be adhered to since, in the original publication, “ The […] use of an inappropriate connecting vowel is not to be considered inadvertent errors ”. Therefore, despite being confusing or misspelled, this name must not be corrected as it is valid in its original spelling; S. oraniensis .

Figures

Figure 1. Representatives of Scolopendromorpha from the Iberian Peninsula (uncollected) A family Plutoniumidae Bollman, 1893: Theatops erytrocephalus (C. L. Koch, 1847), Seoane do Caurel, Lugo (credit: J. Tizón) B family Cryptopidae Kohlrausch, 1881: Cryptops sp. Leach, 1814, Villena, Alacant (credit: D. Molina) C – I family Scolopendridae Leach, 1814 C Scolopendra cingulata Latreille, 1829, Benilloba, Alacant (credit: M. Huesca and CD) and D – I S. oraniensis Lucas, 1840. Observe that in this species the colour variants can occur sympatrically D, H, I Benilloba, Alacant and E – G Almeria Province, Andalucia (credit: F. Rodriguez Luque).

Figure 2. A topotype collection localities in the Iberian Peninsula (southwestern Europe): S. viridipes topotypes (= S. oraniensis); V 1 = Moixent and V 2 = Xàtiva (previously San Felipe), Valencia province, Valencian Country / Community, Spain and S. chlorotes topotypes (= S. oraniensis); C 1 = Málaga municipality, C 2 = Alahurín de la Torre, C 3 = Casabermeja, C 4 = Totalán and C 5 = Estepona, Málaga province, Andalucía, Spain. The “ O ” points S. oraniensis type localities, O 1 = Santon’s mountains [corrected from “ Sauton ” (sic.); Lucas 1846] and O 2 = “ Between Oran and Mers-el-Kebir ”, Oran province (Wilayah), Algeria B “ S. viridipes ” topotype 2, female, (CEUAMr 22) from C Moixent, Alacant D “ S. chlorotes ” topotype 6, female, (CEUAMr 30) from E Alahurín de la Torre, Málaga.

Figure 3. Scolopendra viridipes neotype (= S. oraniensis); male; (“ topotype 1 ", CEUAMr 21) A cephalic plate, antennae, and TT 1–5 B forcipular segment and sternites 1–3 C tooth plates D tergites 13–18 E sternites 10–15. Scale bars: 0.2 mm (C); 1 mm (A, B); 2 mm (D, E).

Figure 4. Scolopendra viridipes neotype (= S. oraniensis); male; (“ topotype 1 ", CEUAMr 21) A ULBS and UL ’ s dorsal view B ULBS ’ s and UL ’ s ventral view C ULBS ’ s, coxopleuron, and UL ’ s ventral lateral view. Scale bars: 1 mm (A, B); 2 mm (C).

Figure 5. Scolopendra chlorotes neotype (= S. oraniensis); male; (“ topotype 5 ", CEUAMr 29) A cephalic plate, antennae, and TT 1–3 B forcipular segment and sternites 1, 2 C tooth plates D tergites 10–14 E sternites 8–12. Scale bars: 0.2 mm (C); 1 mm (A, B); 2 mm (D, E).

Figure 6. Scolopendra chlorotes neotype (= S. oraniensis); male; (“ topotype 5 ", CEUAMr 29) A ULBS ’ s and UL ’ s dorsal view B ULBS ’ s and UL ’ s ventral view C ULBS ’ s, coxopleuron, and UL ’ s ventral lateral view. Scale bars: 1 mm (A, B); 2 mm (C).

Figure 7. Scolopendra oraniensis syntype “ 1 "; unsexed; (MNHN) A cephalic plate, antennae, and T 1 B forcipular segment and sternites 1, 2 C tooth plates D tergites 3–6 E sternites 6–8. Scale bars: 0.2 mm (C); 1 mm (A, B); 2 mm (D, E).

Figure 8. Scolopendra oraniensis syntype “ 1 "; unsexed; (MNHN) A dorsal view of ULBS and UL B ventral view of ULBS and UL C lateral view of ULBS and UL D Original manuscript labels ordered chronologically. Top: old label [Scolopendra canidens Newp. oraniensis Luc. H. Lucas. [Leg.] Oran [type locality]; Middle: Kraepelin’s 1903 labels [“ Type de H. Lucas. Scol. oraniensis Luc. (non canidens Newp.) ”]. Lower: MNHN staff “ modern ” summary label. Scale bars: 1 mm (A, B); 2 mm (C).

Figure 9. Scolopendra oraniensis genitalia A male (CEUAMr 24); and B female (CEUAMr 31), and detail of the setae (arrows) on tarsus 1 (C, D) on the same specimens C male and D female. Observe that on the contrary that Würmli (1980) indicated, the numbers of setae are actually similar in both genders. Legend: AV, anal valve; G. gonopod (folded back); LA, lamina adanalis; LS, lamina subanalis; P, penis; SGS I, sternite of genital segment 1; SGS II, sternite of genital segment 2. Scale bars: 0.1 mm.