Research Article |
Corresponding author: Carles Doménech ( carles.domenech.perez@gmail.com ) Academic editor: Alessandro Minelli
© 2024 Carles Doménech.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Doménech C (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: 49-80. https://doi.org/10.3897/zookeys.1208.122126
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The taxonomic identities of the two largely neglected Scolopendra Linnaeus, 1758 species from continental Spain, S. viridipes Dufour, 1820 and S. chlorotes L. Koch in Rosenhauer, 1856, are examined in this paper. After efforts in locating both species’ type series in eight European institutions, the specimens are considered to be lost. Consequently, the identifications of both taxa were approximated by collating their descriptions with the morphology of all other sympatric Scolopendromorpha. Then, compatible topotypes for both species were collected, and among these a neotype for each taxon were selected and compared with the type series of their respective closest relatives. Finally, both S. viridipes and S. chlorotes are proposed to be conspecific with S. oraniensis Lucas, 1846. Therefore, the name S. viridipes is here established as (senior) syn. nov. and nomen oblitum of S. oraniensis, S. oraniensis is declared as nomen protectum, and S. chlorotes (junior) syn. nov. is reallocated to S. oraniensis. Moreover, the specimens making up the type series of S. oraniensis are also indicated and redescribed, the genitalia are illustrated for the first time, and its specific epithet is briefly reviewed, remaining unaltered in respect of its original spelling.
Chilopoda, chlorotes, continental Spain, oraniensis, Scolopendra, viridipes
The class Chilopoda Latreille, 1817 is a group of venomous terrestrial predators that play an important ecological role in the warm and temperate ecosystems. Taxonomically, this group of arthropods encompasses approximately 3,100 species distributed in five orders (
In peninsular Spain, the order Scolopendromorpha is represented by three of those six families. One of them, Plutoniumidae Bollman, 1893 (Fig.
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).
With the general exception of the north and northwest, these two Scolopendra species are widely and almost sympatrically distributed in continental Spain (
However, besides S. semipedalis Dufour, 1820 (= Himantarium gabrielis (Linnaeus, 1767), see
A topotype collection localities in the Iberian Peninsula (southwestern Europe): S. viridipes topotypes (= S. oraniensis); V1 = Moixent and V2 = Xàtiva (previously San Felipe), Valencia province, Valencian Country/Community, Spain and S. chlorotes topotypes (= S. oraniensis); C1 = Málaga municipality, C2 = Alahurín de la Torre, C3 = Casabermeja, C4= Totalán and C5 = Estepona, Málaga province, Andalucía, Spain. The “O” points S. oraniensis type localities, O1 = Santon’s mountains [corrected from “Sauton” (sic.);
The second species in this underappreciated group is S. chlorotes L. Koch in Rosenhauer, 1856, another small, but much better described taxon from “near Málaga”, Andalusia (south of continental Spain) (Fig.
After more than nine decades of bibliographic silence, the uncertain taxonomic and nomenclatural identities of the two enigmatic species S. viridipes and S. chlorotes are now evaluated, a specific diagnosis for each species is given, and the nomenclatural situation for these taxa is finally clarified.
The depositories of the types series of Scolopendra viridipes, S. chlorotes, and S. oraniensis were scrutinised following the texts of
CLD Cercle Léon Dufour. Saint Severe, Nouvelle-Aquitaine, France
FAU Friedrich-Alexander Universität, Erlangen-Nürnberg, Erlangen, Germany
NHMN Naturhistorisches Museum Nürnberg, Nürnberg, Germany
SAE Sociedad Andaluza de Entomología, Dos Hermanas, Sevilla, Spain
SLB Société Linnéenne de Bordeaux, Bourdeaux, France
UPV Universitat Politècnica de València, València, Spain
Topotypes locations of Scolopendra viridipes are based on information given in
The type series of Scolopendra oraniensis in the
AP apical spines
DM dorso-median spines
LS lateral spines
M median spines
S, SS sternite, sternites
SAP subapical spines
SP prefemoral process spines
T, TT tergite, tergites
UL ultimate legs
ULBS ultimate leg-bearing segment
V ventral spines
VL ventro-lateral spines
VM ventro-median spines
Genital region
AV anal valve
LA lamina adanalis
LS lamina subanalis
SGS I sternite of genital segment 1
SGS II sternite of genital segment 2
The species identifications and differential diagnosis in Tables
The original descriptive morphology of Scolopendra viridipes in comparison to other Scolopendromorpha species described in Peninsular Spain. Bold letters indicate compatible features in other species. * indicates common to the entire Scolopendromorpha, interpretable, or poorly differentiated feature. Nrpp= Not reported, presence possible.
Scolopendra viridipes Dufour, 1820 | S. cingulata Latreille, 1829 | S. oraniensis Lucas, 1846 | Cryptops (Cryptops) anomalans Newport, 1844 | C. (C.) hispanus Brölemann, 1920 | C. (C.) hortensis s.s (Donovan, 1810) | C. (C.) lobatus Verhoeff, 1931 | C. (C.) parisi s.s Brölemann, 1920 | C. (C.) trisulcatus Brölemann, 1902 | C. (Trygonocryptops) longicornis (Ribaut, 1915) | C. (T.) similis Machado, 1953 | Plutonium zwierleini Cavanna, 1881 | Theatops erythrocephalus (C. L. Koch, 1847) | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Presence in Valencianish Kingdom | Present | Present | Present | Nrpp | Present | Nrpp | Absent | Nrpp | Nrpp | Absent | Nrpp | Absent | Present |
Ecology | Edaphic epigean (Lapidicola) | Edaphic epigean, troglophile | Edaphic epigean, troglophile | Edaphic epigean | Edaphic epigean, troglophile* | Edaphic epigean, troglophile* | Edaphic epigean | Edaphic epigean | Edaphic epigean | Troglobiotic | Edaphic epigean | Edaphic, hipogean, troglophile | Edaphic epigean, troglophile |
Colouration | Marked pale, with antenna and legs greenish | Tergites yellowish ended with black distal edge, legs and antenna yellow to orange | Pale cream to dark green or brown with antenna and legs yellowish, greenish, bluish or reddish | Pale yellow to reddish | Pale yellow to reddish | Reddish, sometimes pale yellow | Pale yellow to reddish | Reddish | Pale yellow | Pale yellow | Pale yellow | Pale reddish, legs and antennae usually yellowish | Pale yellow to reddish, cephalic plate and UL usually darker |
Length (in mm) | 40.6 (18 lignes) | 100-155 | 40-65 | 20-40 | 16-30 | 12-30 | 10-13.5 | 14-21 | 15-35 | 27-38 | 30 | 50-80 | 20-45 |
Antennal articles’ number* | More than 15* | 18 | 18-19 | 17 | 17 | 17 | 17 | 17 | 17 | 17 | 17 | 17 | 17 |
Antennae end in a setaceous point* | Yes* | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes |
Headplate shape* | Small* and oval | Oval | Oval | Oval | Oval | Oval | Oval | Oval | Oval | Oval | Oval | Oval | Oval |
Shape of the distal palp’s article (article 3 in second maxillae’s telopodite) * | Dilated and round * | Subcylindrical | Subcylindrical | Subcylindrical | Subcylindrical | Subcylindrical | Subcylindrical | Subcylindrical | Subcylindrical | Subcylindrical | Subcylindrical | Subcylindrical | Subcylindrical |
1st, 2nd and last tergites different in shape respect the others* | Yes* | Yes, smaller in comparison | Yes, smaller in comparison | Yes, smaller in comparison | Yes, smaller in comparison | Yes, smaller in comparison | Yes, smaller in comparison | Yes, smaller in comparison | Yes, smaller in comparison | Yes, smaller in comparison | Yes, smaller in comparison | Yes, 1st and 2nd smaller in comparison, the 21st much bigger | Yes, 1st and 2nd smaller in comparison, the 21st much bigger |
Ultimate legs length respect the locomotory legs | Longer than locomotor ones | Longer than locomotor ones | Longer than locomotor ones | Longer than locomotor ones | Longer than locomotor ones | Longer than locomotor ones | Longer than locomotor ones | Longer than locomotor ones | Longer than locomotor ones | Much longer than locomotor ones | Longer than locomotor ones | Similar in length to locomotor ones, shape markedly thicker | Similar in length to locomotor ones, shape markedly thicker |
The original descriptive morphology of Scolopendra chlorotes in comparison to other Scolopendromorpha species described in Peninsular Spain. Bold letters indicate compatible features in other species. * indicates common to the entire Scolopendromorpha, interpretable, or poorly differentiated feature. Nrpp= Not reported, presence possible. NA= Information not available.
Scolopendra chlorotes L. Koch in Rosenhauer, 1856 | S. cingulata Latreille, 1829 | S. oraniensis Lucas, 1846 | Cryptops (Cryptops) anomalans Newport, 1844 | C. (C.) hispanus Brölemann, 1920 | C. (C.) hortensis s. s. (Donovan, 1810) | C. (C.) lobatus Verhoeff, 1931 | C. (C.) parisi s. s Brölemann, 1920 | C. (C.) trisulcatus Brölemann, 1902 | C. (Trygonocryptops) longicornis (Ribaut, 1915) | C. (T.) similis Machado, 1953 | Plutonium zwierleini Cavanna, 1881 | Theatops erythrocephalus (C. L. Koch, 1847) | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Presence in Malaga | Present | Present | Present | Nrpp | Present | Nrpp | Absent | Nrpp | Nrpp | Present | Present | Present | Present |
Length (in mm) | 42.22 (20 lignes) | 100-155 | 40-65 | 20-40 | 16-30 | 12-30 | 10-13.5 | 14-21 | 15-35 | 27-38 | 30 | 50-80 | 20-45 |
Colouration | Head-plate, antenna and tergites, brownish-green, maxillipeds, last tergite and UL reddish brown, and legs yellowish proximally and greenish distally | Tergites yellowish ended with black distal edge, legs and antenna yellow to orange | Pale cream to dark green or brown with antenna and legs yellowish, greenish, bluish or reddish | Pale yellow to reddish | Pale yellow to reddish | Reddish, sometimes pale yellow | Pale yellow to reddish | Reddish | Pale yellow | Pale yellow | Pale yellow | Pale reddish, legs and antennae usually yellowish | Pale yellow to reddish, cephalic plate and UL usually darker |
Headplate shape* | Oval | Oval | Oval | Oval | Oval | Oval | Oval | Oval | Oval | Oval | Oval | Oval | Oval |
Tooth-plate on coxosternum | Present | Present | Present | Absent | Absent | Absent | Absent | Absent | Absent | Absent | Absent | Present | Present |
Tergites shape* | Tergite sides straight, TT1-3 and 20 rounded anteriorly | Quadrangular TT1-2 and T21 slightly smaller | Quadrangular TT1-2 and T21 slightly smaller | Quadrangular TT1-2 and T21 slightly smaller | Quadrangular TT1-2 and T21 slightly smaller | Quadrangular TT1-2 and T21 slightly smaller | Quadrangular TT1-2 and T21 slightly smaller | Quadrangular TT1-2 and T21 slightly smaller | Quadrangular TT1-2 and T21 slightly smaller | Quadrangular TT1-2 and T21 slightly smaller | Quadrangular TT1-2 and T21 slightly smaller | Quadrangular TT1-2 slightly smaller, Segment 21 longitudinal enlarged | Quadrangular TT1-2 slightly smaller, Segment 21 longitudinal enlarged |
TT’s paramedian sutures | TT2-20 | TT2/3-20 | TT2-20 | TT2-20 | Sutures weak, apparently present since T2 | TT3, 4 or 5-20 | TT2-20 | TT2/3-20 | TT2-20 | TT2, 3 or 4-20 | TT2-20 | TT2-20 | TT2-20 |
Tergites margination* | TT14-21 | TT7/12-21 | Complete from TT19-21 | TT3-19 | NA | TT3, 4 or 5-20* | NA | NA | TT3-20 | TT4-21 | TT3-20 | T21 | T21 |
Median longitudinal suture on the T21 | Present | Absent | Present | Presence observed variable; T21 with a depression | NA | NA, probably absent; T21 with a depression | NA, probably absent | Absent | Present | Absent | NA | Present | Present |
Paramedian sutures on sternites | Longitudinal, absent in S 21 | SS2-20 | SS2-20 | Cruciform sutures | Cruciform sutures | Cruciform sutures | Cruciform sutures | Cruciform sutures | Cruciform sutures | Trigonal sutures | Trigonal sutures | Single medial longitudinal suture | Single medial longitudinal suture |
UL’s prefemoral process | Short, with two blunt spines at the extreme | Short, with 3-5 spines | Short, with 2 or 3, rarely up to 6 spines | Absent | Absent (femur and tibia with 4 distal processes) | Absent | Absent | Absent | Present, with a single spine (femur with a distal process) | Absent | Present, with a single spine (femur with a distal process, tibia with distal two processes) | Process absent | Process absent |
UL’s spinous prefemoral formula | V: 19 spines arranged in for rows, M:7 spines | VL:0; V: 2; VM: 0; M: (1)2; DM: (1)2 | VL:3-5; V: 3-6; VM: 2-6; M: 4-8; DM:2-4 | Prefemur with setae, spines absent,7-10 saw teeth on tibia and 3-5 on tarsus 1 | Prefemur with setae, spines absent, 8 saw teeth on tibia and 4 on tarsus 1 | Prefemur with setae, spines absent, 5-8(9) saw teeth on tibia and 2-4 on tarsus 1 | Prefemur with setae, spines absent, 6-10 saw teeth on tibia and 4-6 on tarsus 1 | Prefemur with setae, spines absent, 6-12 saw teeth on tibia and 4-8 on tarsus 1 | Prefemur with setae, spines absent, 13 saw teeth on tibia and 4 on tarsus 1 | Prefemur with setae, spines absent, 12-13 saw teeth on tibia and 5 on tarsus 1 | Prefemur with setae, spines absent, 10 saw teeth on tibia and 3 on tarsus 1 | Spines absent | With one ventromedial prefemoral and one femoral spine |
The distribution of Scolopendra species colour variants were defined using the BV (
Background removal in illustrations, contrast adjustments, map configuration, and their respective clarifying notes were performed with Adobe Photoshop CS6 software®. A base map was obtained from the National Oceanic and Atmospheric Administration, National Weather Service (
Scolopendra oraniensis type series morphological comparison with S. viridipes’ and S. chlorotes’ topotypes (= S. oraniensis; composite data from Supplementary tables 1 and 2). AP, apical spines; SAP, subapical spines; DS, dorsal spines; LS, lateral spines; VL, ventro-lateral spines; V, ventral spines; VM, ventro-median spines; M, median spines; DM, dorso-median spiness; SP, prefemoral process spines; UL; ultimate legs, ULBS, ultimate leg-bearing segment; T, tergite; TT, tergites, S, sternite; SS, sternites, RG, Retracted genitalia; * No visible (cephalic plate flexed over firsts SS), interpretable or damaged.
Scolopendra oraniensis (type series) | S. viridipes (CEUAMr21-25) | S. chlorotes (CEUAMr26-40) | |||||||
---|---|---|---|---|---|---|---|---|---|
Syntype “1" | Syntype “2" | Syntype “3" | Syntype “4" | Syntype “5" | Syntype “6" | Syntype “7" | |||
Body length in mm | 64 | 60 | 58 | 52 | 42 | 41 | 41 | 35-40 | 26-60 |
Sex | RG | RG | RG, probably female | RG, probably female | RG | RG | RG | – | – |
Antenna reaching to tergite* | T2* | T2* | T2* | T2* | T2* | T2* | T2* | T3 | T3 |
Number of antennal articles | 18/19 | 18/19 | 19/18 | 19/19 | 19/18 | 19/19 | 18/19 | 17-20 | 17-20 |
Number of proximal glabrous articles | 5½ | 5½ | 5½ | 5½ | 5½ | 5½ | 5½ | 5 | 5-5½ |
Teeth on tooth plate | 4+4 | * | 4+4 | 4+4 | 3+3 | 4+4 | 4+4 | 4+4 | 3+3; 4+4 |
Teeth on forcipular trochanteroprefemoral processes Total (apical /medial) | 3(1/2) - 3(1/2) | * | 3 (1/2) - 3 (1/2) | 2 (1/1) - 3 (1/2) | 2 (1/1) - 2 (1/1) | 3(1/2) - 3(1/2) | 3(1/2) - 3(1/2) | 3(1/2) | 2(1/1); 3(1/2) |
Tergite paramedian sutures | TT2-20 | TT2-20 | TT2-20 | TT2-20 | TT2-20 | TT2-20 | TT2-20 | TT2-20 | TT2-20 |
Longitudinal suture on T21 | Present | Present | Present | Present | Present | Present | Present | Present | Present |
First tergite with complete margination | 19 | 19 | 19 | 19 | 19 | 19 | 19 | 17-19 | 14-18 |
Paramedian sutures on sternites | SS2-20 | SS2-20 | SS2-20 | SS2-20 | SS2-20 | SS2-20 | SS2-20 | SS2-20 | SS2-20 |
Spines in coxopleuron (Left/Right) | 1/1 | 1/1 | 1/1 | 1/1 | 1/1 | 1/1 | 1/1 | 1/1 | 1/1 |
Spines in coxopleural process (Left/Right) | AP-SAP: 4/5 DS: 2/1 LS: 4/4 Total:10/10 | AP-SAP: 5/4 DS: 2/1 LS: 4/4 Total:11/9 | AP-SAP: 6/5 DS: 1/2 LS: 6/6 Total:13/13 | AP-SAP: 5/6 DS: 3/2 LS: 6/4 Total:14/12 | AP-SAP: 4/5 DS: 2/2 LS: 4/6 Total:10/12 | AP-SAP: 6/5 DS: 3/2 LS: 3/4 Total:12/11 | AP-SAP: 5/5 DS: 3/3 LS: 4/5 Total:12/13 | AP-SAP: 4-8 DS: 1-3 LS: 2-3 Total: 7-14 | AP-SAP: 4-7 DS: 0-4 LS: 1-4 Total: 5-15 |
Tarsal spurs on leg 1 (Left/Right) | 2/2 | 2/2 | 2/1 | 2/2 | 2/2 | 2/2 | 2/2 | 0-2 | 1-2 |
Legs with one tarsal spur | 2-19 (Left 1-18) | 2-19 | 2-19 | 2-19 (Right 1-18) | 2-19 | 2-19 | 2-19 | 2-19 | 2-18 or 19 |
Ultimate legs prefemoral spinulation and spines in prefemoral process (Left/Right) | VL:4/3 V: 6/6 VM: 4/4; M: 4/4; DM:2/2; SP:2/2; Total: 22/21 | VL: 3/5; V: 3/5; VM: 5/6; M: 8/6; DM:4/4; SP:3/2; Total: 26/28 | VL:1/3; V: 3/2; VM: 4/4; M: 5/5; DM:2/2; SP:3/4; Total: 18/20 | VL:5/5; V: 5/4; VM: 3/2; M: 5/6; DM:2/2; SP:3/3; Total: 23/22 | VL:4/5; V: 4/6; VM: 5/4; M: 4/6; DM:3/4; SP: 2/3; Total: 22/21 | VL:6/5; V: 5/5; VM: 3/5; M: 3/6; DM:2/2; SP:2/2; Total: 21/25 | VL: 3/5; V: 3/5; VM: 5/6; M: 6/6; DM:2/3; SP:3/2; Total: 22/27 | VL: 3-7; V: 4-6; VM: 3-7; M: 5-8; DM: 2; SP: 2-3; Total: 19-33 | VL: 2-10; V: 4-10; VM: 2-8; M: 2-10; DM: 1-7; SP: 0-5; Total: 11-50 |
Scolopendra oraniensis syntype “1"; unsexed; (
Order Scolopendromorpha Pocock, 1895
Family Scolopendridae Leach, 1814
Subfamily Scolopendrinae Kraepelin, 1903
Tribe Scolopendrini Leach, 1814
Genus Scolopendra Linnaeus, 1758
Scolopendra viridipes Dufour, 1820: 317.
?S. clavipes C. L. Koch, 1836 [1847 sic.]:
S. viridipes: Walckenaer and Gervais 1841: 258, as unrecognisable taxon.
?S. doriae Pirotta, 1878a (= S. cingulata):
?S. oraniensis
S. viridipes
[Dufour, 1822 sic.]:
From Latin viridis (green) and pes (feet), literally meaning green-footed Scolopendra.
Types currently lost.
J. M. L. Dufour, between 1811 and 1813 (
“Kingdom of Valencia”, Valencian Community, east of Spain.
As for type locality.
With the express purpose of clarifying the taxonomic status and the type locality of the nominal taxon S. viridipes Dufour, 1820, the following neotype for this species is designated (
Male. Embassament del Bosquet, Moixent; Valencia Province (Spain) (38°51'21.7"N, 0°44'36.7"W 380 m a.s.l.) (Figs
S. viridipes Dufour, 1820 is an invalid name subjectively designated here as nomen oblitum and a senior synonym of S. oraniensis Lucas, 1846.
[annotations in brackets]. (Original description available from: https://books.google.be/books?vid=GENT900000003803&printsec=frontcover#v=onepage&q&f=false)
XII. Green-footed Scolopendra
Scolopendra (viridipes) shell [dorsal habitus; tergites] livid, antennae and feet [legs] greenish, 21 feet [legs] in each side, the posterior ones longer.
Habitat: under the rocks of the Valencian Kingdom’s mountains. Length 18 lignes [1 Paris ligne = 2.2556 mm; 40.6 mm].
It differs from Scol. morsitans. The body segments [sensu tergites] are roughly square and equal between them, except for the first two and the last one. The head [cephalic plate] is small and oval. The whole body [dorsal habitus] has a markedly livid colour. The legs and the antennae are greenish. They [antennae] end in a setaceous point and have more than fifteen articles. The legs grow from the head [cephalic plate] to the anus [ultimate leg-bearing segment; ULBS]. The palps [first maxillary telopodites] end with a dilated and round article [article 3].
Efforts to locate S. viridipes type series in its four most probable repositories, CLD,
Knowing that Dufour mostly preserved his specimens in dry conditions (
Therefore, the S. viridipes type series is here considered as definitively lost material.
Because the type material for S. viridipes has vanished, and to determine to which species
Initially, the brief morphological description does not allow one to assign S. viridipes to any definite genus because some features are common for all Scolopendromorpha, either because they are insufficiently detailed or widely observed (See “*” in Table
The six remaining Cryptops species (Fig.
Otherwise the exclusive colouration of S. viridipes, a “livid” dorsal habitus with greenish legs and antennae (Figs
Scolopendra cingulata always has tergites with posterior transverse black pigmentation combined with shiny red, orange, or yellow colouration in their legs when juvenile (
The remaining Scolopendromorpha species inhabiting the presumed type locality of S. viridipes is S. oraniensis. This species is variable in colouration even within local populations (Fig.
Hence, prior to analysing some compatible topotypic material, all features stated in the original description strongly suggest that the closest relative to S. viridipes is S. oraniensis, if they do not belong to the same taxon.
Four colour-, morphological-, and size-compatible S. viridipes topotypes were collected in two Valencian localities known by their authorities (Figs
As the two taxa, S. viridipes Dufour, 1820 and S. oraniensis Lucas, 1846, are deemed to be conspecific, the “Principle of Priority” provides preference for the name S. viridipes to replace the name S. oraniensis (
This statement is not strictly true, since
Hence, the name S. viridipes is here declared invalid since this is subjectively considered a senior synonym and nomen oblitum of S. oraniensis, while the name S. oraniensis is proposed as a nomen protectum, being fixed for unequivocal referencing of this species (
Scolopendra chlorotes L. Koch in Rosenhauer, 1856: 417.
?S. oraniensis Lucas, 1846:
S. chlorotes:
From the Greek word χλοερός (khloerós, verdant) and χλόη (khlóē, “the green of new growth”) meaning greenish yellow, pale green, pale, pallid, or verdant, referencing their pale greenish and yellowish leg colouration.
Types currently lost.
W. G. Rosenhauer, in 1849 (
“Near Málaga”, Andalusia, Spain.
As for type locality.
With the express purpose of clarifying the taxonomic status and the type locality of the nominal taxon S. chlorotes L. Koch in Rosenhauer, 1856, the following neotype is designated (
Male. Carretera de Coín, Alahurín de la Torre, Málaga province (Spain) (36°39'37'’N, 4°31'0.4'’W 104 m a.s.l.) (Figs
Scolopendra chlorotes L. Koch in Rosenhauer, 1856 is an invalid name, here subjectively proposed as a junior synonym of Scolopendra oraniensis Lucas, 1846.
[annotations in brackets]. (Original description available from: https://www.biodiversitylibrary.org/page/42185817#page/425/mode/1up)
Scolopendra chlorotes Koch.
Brownish green, feeding pliers [forcipules], end-shield [ULBS tergite] and last pair of legs [terminal legs] reddish brown, on the last seven tergites, a furrow at the lateral edges [margination in tergites], on the first podomere of the last pair of legs [UL prefemur] 19 small teeth on the underside [ventral position], seven small teeth directed inward on the upper-side [medial and dorsomedial positions].
Length 20 ‘lignes’ [1 German – Nuremberg – lignes ≈ 2.11 mm; 42.22 mm].
Shiny; head [cephalic plate] longish, rather narrow, the head area dentate [probably referring to tooth plates] in the middle, tergite sides straight, those of the 3rd and the penultimate tergite rounded anteriorly, the seven last tergites with a furrow at the lateral edges [tergite margination]. Tergites with the two normal stripes [paramedian sutures], except the first and the last tergites; the end-shield [ULBS tergite] shows a distinct median longitudinal furrow [suture]. The sternites have two longitudinal furrows [paramedian sutures], except the last sternite. The last pair of legs short, dorsally flat, the tooth-like process at the inner angle of the first podomere short [UL prefemoral process] with two blunt teeth [spines] at the tip; at the inner side [medial position] of this podomere [UL prefemur] seven small teeth [spines] at the upper side [medial position], 19 small teeth [spines] arranged in four rows at the underside [ventral position]. Head [cephalic plate], antennae, and tergites, except the last one, brownish green, the first podomeres [prefemur, femur, and even tibia] of the legs yellowish, the last [tibia and tarsi 1 and 2] green; maxillipeds, lower lip [probably trochanterprefemoral parts of the forcipula, tooth plates or/and coxosternite], end-shield [ULBS tergite] and last pair of legs [UL] reddish brown, the capture-claw [forcipules] brownish black from the middle [tarsungula].
Near Malaga, sporadic.
All attempts to locate the type series of S. chlorotes were unsuccessful; the curators in their four probable depositories (NHMN,
According to
The other smaller part of Koch’s wet collection, probably containing this species type series, did remain at the NHMN. However, the NHG rooms, their catalogues, and a large part of the collections were damaged during World War II in 1945 and presumably also the S. chlorotes types (EM Neupert (NHG) pers. comm. Sep. 2020). Therefore, the type series of S. chlorotes is considered to be lost.
Because the type material is unavailable, the features in the original description are compared to those of other Scolopendromorpha found in Peninsular Spain to determine to which taxon L. Koch was referring when he erected S. chlorotes as a new species (Table
Hence, its unambiguous morphology and colouration, compatible location, and the exclusion of all other Iberian Scolopendromorpha are facts that, combined, strongly suggest that if it is not the same taxon, the closest relative to S. chlorotes is S. oraniensis (Table
A total of fifteen S. chlorotes topotypical specimens were collected in five municipalities “near Málaga” (L.
Additionally, the neotype of S. chlorotes and the other “topotypes” were compared with those of S. viridipes, and conspecificity was also confirmed (compare Figs
As long as the taxon S. chlorotes L. Koch in Rosenhauer, 1856 is recognised as conspecific with S. oraniensis Lucas, 1846, the principle of priority indicates that the valid name of S. chlorotes is S. oraniensis (
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:
S. canidens lusitanica Verhoeff, 1931: 309.
S. oraniensis:
[based on S. oraniensis type series]. Body length up to 64 mm. 18 or 19 antennal articles; 5½ 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. T1 without sutures or sulci. Paramedian sutures complete on tergites TT2–20. T21 with a complete median longitudinal suture. Margination starting at T14, complete on TT19–21. Sternite paramedian sutures on TT2–20. 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/T21 length ratio ≈ 5.15.
from the toponym “Oran”, Algeria and the feminine (or masculine) suffix -ensis (from) meaning “from Oran”, Algeria, in reference to the type locality.
P. H. Lucas, during winter, between 1839 and 1842 (
from
Known from southern France (including Corsica), southern Italy (including Sardinia and Sicilia), Malta, Spain (including Balearic Islands), Portugal, Morocco, and Algeria. Introduced in Japan (
Nomen protectum.
(Table
(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 [...].
[notes on brackets are comprehensive annotations from this text author]: Body length up to 64 mm. Antennae reaching up to T2 [maybe up to T3; actual length shrivelled by ethanol retraction]; with 18 or 19 articles, the basal 5½ ones dorsally and ventrally glabrous (Fig.
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 T1 (Fig.
Spiracles triangular with three valves, present on body segments 3, 5, 8, 10, 12, 14, 16, 18, and 20.
T1 without sutures or sulcus (Fig.
Sternites with complete paramedian sutures from TT2–20 (Fig.
Coxopleuron not surpassing the posterior border of the tergite of the ULBS (Fig.
Leg 1 with two distal tarsal spurs on tarsus 1, one lateral anterior and one ventral; legs 1–18 or 19 (mode 19, Table
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.
Genitalia retracted in the whole type series.
(Based on S. viridipes and S. chlorotes “topotypical” material; Fig.
Scolopendra oraniensis genitalia A male (CEUAMr24); and B female (CEUAMr31), 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
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.
Scolopendra oraniensis has been observed to be somewhat variable in some few morphological features (Figs
Additionally, the colouration has proven to be variable sympatrically (Figs
These morphological and colouration characteristics, although tending towards a certain geographic distribution, often turn out to be variable within those populations (Table
Morphologically, S. oraniensis is related to five other species of the S. canidens group distributed around the Mediterranean Sea and Middle Asia (
In these last two species closest to S. oraniensis, comparative analyses such as antennae/cephalic plates and T21/UL length ratios and their genitalia descriptions have not been performed yet.
To verify the actual identities of S. viridipes and S. chlorotes, establishing the S. oraniensis specimens
At a nomenclatural level, the specific epithet of S. oraniensis obviously conformed to the reference of the type locality (
The identification of species that were described during the early time of binomial nomenclature (18th and 19th centuries) can often be a complicated task. The usually short and superficial descriptions as well as the frequent disappearance of type series are among the main reasons hindering these tasks. In that time, the limited access to information and collections, and the insufficient faunistic and taxonomic knowledge frequently caused the ignorance of some species names, or, on the contrary, the unintentional creation of many synonyms for the same taxon, with authors working individually in different countries with little or no communication. Since their description, the combination of these scenarios has affected the identities of S. viridipes and S. chlorotes.
After concluding on these species conspecificity with S. oraniensis, it can be observed that Lucas, when he described in 1846 his “greenish legged Scolopendra”, he accidentally overlooked the legs colour similarity between his “new” species and
So far, the incorporation of molecular data has provided interesting insights in to the systematics of Scolopendra (
Even in the absence of modern methodologies and since the present contribution has proven useful, other authors are encouraged to continue producing studies with this classic taxonomic approach. These endeavours could help to improve the current Chilopoda knowledge by clarifying the identity of some taxa described long ago, as it has been the case of S. viridipes, S. oraniensis, and S. chlorotes.
The author sincerely thanks S. Rojo (UA), V. M. Barberá (UA) and E. Larriba (UMH; Universitat Miguel Hernández, Elx, Spain) for their constructive comments on the manuscript. Also, thanks are extended to P. Gaboriaud (CLD), J. J. Geoffroy and E. Leguin (
Finally, CD wishes to express his heartfelt gratitude to D. Cabanillas (UM; Universidad de Murcia, Spain) for his important and unselfish assistance, which has undoubtedly been critical to the completion of this work.
The author has declared that no competing interests exist.
No ethical statement was reported.
No funding was reported.
The author solely contributed to this work.
Carles Doménech https://orcid.org/0000-0003-1890-9434
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Supplementary information
Data type: docx
Explanation note: file 1. Scolopendra viridipes (=S. oraniensis) neotype and “topotypes” morphological comparisons. AP, apical spines. SAP, subapical spines. DS, dorsal spines. LS, lateral spines. VL, ventro-lateral spines. V, ventral spines. VM, ventro-median spines. M, median spines. DM, dorso-median spines. SP, prefemoral process spines. UL, ultimate legs. ULBS, ultimate leg-bearing segment. T, tergite. TT, tergites. S, sternite. SS, sternites. RG=Retracted genitalia. *= Not visible, damaged, or regenerated. file 2. List of references satisfying the second requisite of reversal precedence of the principle of priority (