Scolopendra aztecorum Verhoeff, 1934

Martínez-Muñoz, Carlos A., Dolejš, Petr, Kronmüller, Christian, D-, ., ., -, ., ., D- & ., 2016, On the true identity of Scolopendra aztecorum Verhoeff, 1934 (Chilopoda: Scolopendromorpha: Scolopendridae), Ecologica Montenegrina 7, pp. 405-413 : 406-411

Scolopendra aztecorum Verhoeff, 1934

( Figs. 1 , 2a–h )

Scolopendra azteka Verhoeff , in litt. (non Scolopendra azteca Saussure, 1858 ) (See remarks) Scolopendra aztekorum Verhoeff, 1934: 49 . (See remarks)

Scolopendra aztecorum Verhoeff, 1934: 49–50 .

Scolopendra aztecorum Bücherl, 1942: 291 , 292, 293.

Scolopendra aztekorum Moritz & Fischer, 1979: 329 .

Scolopendra aztecorum Shelley, 2006: 27 , 40.

Scolopendra aztecorum Minelli et al., 2006 –2015.

Scolopendra aztecorum Cupul-Magaña, 2009: 51 .

Scolopendra aztecorum Cupul-Magaña, 2013: 35 .

Scolopendra aztecorum Dolejš in Tuf & Tajovský, 2014: 20.

Scolopendra aztecorum Dolejš in Tuf & Tajovský, 2015: 7.

Scolopendra aztecorum Bonato et al., 2016 .

Type material. Verhoeff (1934) described S. aztecorum based on six syntypes.

Type locality. Mexico, Baja California Sur , near La Paz and near Los Inocentes ( Fig. 1 ). See remarks. Distribution. Known only from type localities. Endemic to Mexico .

Original rank. Species.

Current rank and status. Valid species.

Material examined for redescription. Five syntypes. NMP P6 E-1303, one adult specimen. | Scolopendra aztecorum VERHOEFF, 1937 . det. L. J. Dobroruka. La Paz , J. Kalifornie, U.S.A. (pod hnijícími kaktusy). 1 kus (SYNTYPUS). lgt. Prof. Dr. K. Štorkán. 1930. Coll. Nár. muzeum. I. č. 1303. P. č. 45/1975. | ZMB 13378 , one juvenile specimen. | Syn-Typus | | Scolopendra azteca Verh. mexican. Kalifornien. 1095 | RMNH.CHIL.154, one juvenile specimen. | Cat. Nº 154 | | Scolopendra azteca Verh. Mexic. Kaliforn. | ZSM- A- 20051044, one juvenile, dissected and in poor condition, all dissected parts are included in the vial, apparently there are no slides by Verhoeff. | Holotype | | Scolopendra aztekorūm Verh. Mexikanisch Californien. | ZSM-A- 20051045, one adult in good condition. | Scolopendra aztekorūm Verhoeff mexik. Californien. Coll. Verhoeff. |

Diagnosis. With shared characters of S. viridis , S. polymorpha and S. chlora Chamberlin, 1942: 1 ) cephalic plate without paramedian sutures, 2) transverse sulcus on tergite 1, 3) two paramedian sutures on tergite 1, 4) median longitudinal suture on tergite 21 and absence of prefemoral spines on legs 19–20. Readily distinguishable from the other three species by the unique character of two spurs on tarsi 1 of legs 1–19(20) against two spurs only present on tarsi 1 of legs 1.

Redescription

Color: According to Verhoeff (1934), body and legs pale yellow, antennae, head and T1 dark bluish, sometimes also tergites 2 and 3, forcipular coxosternite and forcipules reddish-brown, tarsungula black. Only one adult with head and T1 more yellowish-brown. At present, former bluish color on the head and T1 is faded.

Antennae: 27–33 antennal articles, 15–17 basal sparsely hirsute. The transition between sparsely hirsute and hirsute antennal articles is gradual and difficult to see. This may be due to the setae being very short and pale. Setae not arranged in longitudinal rows. ZSM-A- 20051044 with antennae broken, only 15 left and 26 right articles remaining. Bücherl (1942) incorrectly transcribed the count from Verhoeff (1934) as only “27– 32 artículos antenais”.

Cephalic plate: Slightly and sparsely punctate. Short anterior sulcus. Without paramedian sutures. Posterior margin almost straight, overlapping anterior margin of T1.

Mandibles and maxillary complex: Specimen ZSM-A- 20051044 with right mandible, 1st maxillae and 2nd maxillae dissected. Right mandible with five tricuspid teeth, condyle of lamina condilifera well developed. 2nd maxillae articles not ridged or pointed ventrally, article 2 with a lateral spur. According to Verhoeff (1934), the spur (“Dorn”) may be present or absent, but non dissected specimens were not checked for this character to avoid manipulation damage. Article 3 with conspicuous dorsal brush. Claw with 2 accessory spurs.

Forcipular segment: Coxosternite with a short anteromedian longitudinal suture (aprox. 1/8 to less than 1/3 of the coxosternite’s length) (in specimen ZSM-A- 20051045 double and reticulate in the middle), without transversal suture. Tooth-plates wider than long, slightly separated. Typical single short strong seta in the middle of tooth-plates, not seen by Verhoeff (1934): “Die Zahnplatten innen mit Grube, aber ohne borstentragenden tuberkel”. Tooth- plates with 1+1+2 teeth, the lateral tooth more isolated and sharp, the median ones more blunt. Process of trochanteroprefemur large, generally with 2 tubercles on its internal margin (in specimen ZSM-A- 20051045 left process without noticeable tubercles, right with two insinuated bumps, in specimen ZSM-A- 20051044 tubercles well developed). Trochanteroprefemoral process surpassing the anterior border of the tooth-plates. Interior surface of tarsungula with a single sharp longitudinal ridge, smooth, not serrate.

Tergites: T1 with curved anterior transverse sulcus. Two faint paramedian sutures, diverging and branching anteriorly, not seen by Verhoeff (1934). Branching sutures 3 (2–4) each side and sometimes 1 medial, often more sclerotized and darker where they touch the transverse sulcus. The sutures cross the transverse sulcus and form a reticulate pattern anteriorly. TT2–20 with complete paramedian sutures (except specimen ZSM- A- 20051045 with sutures of T3 incomplete in the middle). Paramedian sutures in T2 and T4 anteriorly convergent, in T3 anteriorly divergent. T21 with median longitudinal suture, from complete to incomplete anteriorly (1/10) and posteriorly (3/10). Oblique sutures generally present in T7, T8, T10, T12, T14 and T16. In tergites anterior to T7 there is not a definite oblique suture but a reticulum. Margination starting from T7– T11 to T21 (in ZMB 13378 seen only from T16 to T21), in anterior tergites incomplete and less marked, generally marked and complete from T8–T14 onwards. T21 as wide as long or slightly wider than long, posterior margin rounded.

Sternites: SS2–20 with two complete paramedian sutures. S21 generally longer than wide (but wider than long in NMP P6E-1303), posterior margin straight to emarginate.

Legs: Tarsi 1 longer than tarsi 2. Legs 1–20 without spines. Leg pair 1 with 1 prefemoral, 1 femoral, 1 tibial, 2 tarsal (on tarsi 1) and two pretarsal accessory spurs. Legs 2–20 without tibial spurs, legs 2–19(20) with two tarsal spurs on tarsi 1, sometimes only one tarsal spur on leg pair 20, legs 2–20 with pretarsal accessory spurs.

Coxopleura: Coxopleural surface without setae. Inner half of the coxopleuron not notably warped, in contrast to species like S. angulata Newport, 1844 and S. gigantea Linnaeus, 1758 . Extensive pore-field, covering most of the ventral surface of the coxopleuron, including the posterior margin, the inner margin and part of the coxopleural process, but leaving a short and narrow poreless stripe anterior to the coxopleural process. Coxal pores numerous, small, grouped (generally separated by not more than 1–2 pore diameters). Posterior margin of coxopleuron slightly oblique, with one side spine. Coxopleural process short, conical, with 4–7 spines, from which two are apical and the rest subdistal in ventral, lateral or dorsal position, the generally 4 (3–5) biggest spines organized in a row. Verhoeff (1934) considered the coxopleural process as strongly developed and slender, moderately long, with 3–5 outer spines.

Ultimate legs: Prefemur and femur somewhat flattened dorsally, other articles cylindrical. Pretarsal accessory spurs present, without other spurs. Prefemur dorsodistally barely notched. Only prefemur spined. Prefemoral spines 19–27 (21–24 in Verhoeff, 1934) organized in 5–7 rows (1 dorsolateral internal, 0–1 lateral internal, 0–2 ventrolateral internal, 1–2 ventral, 1–2 ventrolateral external, 0–1 lateral external). Internal dorsolateral row with 4–7 spines. Prefemoral process with 3–5 spines (only 4 in Verhoeff, 1934), in ZSM-A- 20051044 left leg prefemoral process with 5 big and 1 small spine, in NMP P6E-1303 right leg process regenerated and without spines. Prefemur with basal ventral depression, flanked anteriorly by 1–3 small spines.

Size: 55–120 mm. Size as Verhoeff (1934): Adults from 118–125 mm, juveniles from 70–77 mm (see remarks).

Discussion. The posterior margin of the head overlapping T1, T1 with a transverse sulcus, and the distribution of tarsal spurs allow the unequivocal differentiation from A. pygmaeus . The character of two tarsal spurs on legs 1–19(20) allows the recognition of Scolopendra aztecorum as a valid species, different from S. viridis , S. polymorpha , S. chlora or any other New World Scolopendra species. The only known exception is the mistake in the description of the neotype of Scolopendra pomacea by Cupul-Magaña et al. (2015), where the authors wrote “ Scolopendra pomacea differs from the other described species of the genus Scolopendra in Mexico in the following combination of characteristics ( Attems 1930): …tibia of legs 1–19 with one spur and tarsus 1 with two spurs; legs 20 with tibia usually without spur...”. This is clearly an erroneous translation of Attems (1930) description, the correct translation being: “…tarsi 1 of legs 1–19 with one spur and pretarsi with two accessory spurs; legs 20 with tarsus 1 usually without spur...”.

Remarks

Regarding an avoided potential homonymy. Verhoeff (1934) stated in a footnote “ aztekorum = azteka Verh. i. litt. (non azteka Sauss. !)”. This means that he first used the name S. azteka for the species in a letter or other documented correspondence, probably with a date. The existence of this letter is unknown, and the document is not published in the sense of the ICZN. Verhoeff (1934) noted that this name will fall in homonymy with S. azteka (sic!) Saussure, so he changed it to S. aztekorum .

Regarding species epithets. Verhoeff (1934) used “k” instead of “c” in the first mention of the epithet “ aztekorum n. sp. ” in his key for Scolopendra species from the viridis group. He immediately remitted to the aforementioned footnote, and there also used “k” for the epithets aztekorum (it was the second time) and azteka (twice, Verh. and Sauss.). However, he used the binomen Scolopendra aztecorum n. sp. in the species description, immediately after the key and on the same page as the footnote. Verhoeff (1934) utilized the epithet aztecorum two more times, in the description of Scolopendra viridis storkáni [recte storkani ( Shelley, 2006)]. All subsequent authors except Moritz & Fischer (1979) have used the spelling Scolopendra aztecorum . The most frequently cited Scolopendra aztecorum Verhoeff, 1934 is here selected as the correct original spelling. The Scolopendra aztekorum spelling is incorrect (Art 24.2.3, ICZN) and unavailable (Art. 32.4, ICZN). We are the First Revisers (Art. 24.2.3, ICZN).

Regarding type specimen status. Verhoeff (1934) described Scolopendra aztecorum based on six specimens without selecting a holotype, which makes them syntypes (Art. 72.1.1, ICZN). Shelley (2006), followed by Cupul-Magaña (2013), incorrectly stated that the specimen at ZMB was the Scolopendra aztecorum holotype and immediately after cited Moritz & Fischer (1979). However, Moritz & Fischer (1979) cited ZMB specimen as a “ Syntypus ” of Scolopendra aztekorum . This confusion was addressed by Dolejš (2015). It is worth noticing that the specimen ZSM-A- 20051044 is incorrectly labeled as holotype and that the specimens ZSM-A- 20051045 and RMNH.CHIL.154 were not labeled as types.

Regarding the designation of a lectotype. Lectotype designations made after 1999 should have the purpose of clarifying the application of the name to a taxon ( ICZN, 1999, p. 11). This is not the case of Scolopendra aztecorum , so it is not necessary. On the other hand, the type series is shared among at least four institutions in three countries. To declare a lectotype will deprive all institutions but one from the possession of name bearing specimens and it will also affect published references on syntype holding. The loss or destruction of the lectotype will lead to a new nomenclatural act designating a neotype while the loss of one or few syntypes from this series will require no action. Keeping the syntypes avoids two potential nomenclatural acts and maintains stability.

Regarding the missing syntype. The sixth syntype was not found. It is not at any of the BMNH (Greg Edgecombe, pers. comm., 18.xi.2014), NHMW (Nesrine Akkari, pers. comm., 11.xii.2014), ZMUC (Henrik Enghoff, pers. comm., 20.x.2015), MHNG (John Hollier, pers. comm., 1.xi.2015) or NHRS (Karin S. Kronestedt, pers. comm., 21.i.2016). It is listed neither by Weidner (1960) nor by Rack (1974) as being at ZMUH, nor by Stoev & Beron (2000) as being at NMNHS. It is not found in the dataset of the Myriapoda and Onychophora collection of the MNHP cited by Le Bras et al. (2015). Verhoeff’s type series usually consist of wet material and micropreparations of dissected parts. There might be a chance that the sixth syntype is dissected and deposited as slides in one of the European collections that stores part of Verhoeff’s collection.

Regarding type locality. Shelley (2006), Minelli et al. (2006 –2015), Cupul-Magaña (2013) and Bonato et al. (2016) stated that the type locality of S. aztecorum was “ Mexico, Baja California Sur, La Paz”. The recent additions and corrections to Cupul-Magaña (2013) by Flores-Guerrero et al. (2015) did not include new information about the type locality. According to Verhoeff (1934): “4 Erwachsene und 2 Adolescentes stammen vom südlichsten Teil der Halbinsel Kalifornien, wo sie sich unter modernden Kakteenstämmen aufhielten, teils bei La Paz, teils bei Inocentez”. The syntypes were collected near La Paz and near Los Inocentes, not “Inocentez” as mispelled by Verhoeff (1934) and Bücherl (1942). Art. 73.2.3 ( ICZN, 1999) establishes that the type locality encompasses all of the places of origin.

Regarding distribution. Distribution was given as “ type locality”, meaning La Paz only, by Shelley (2006) and “BCS: La Paz” by Cupul- Magaña (2013). Distribution is type locality, La Paz and Los Inocentes.

Regarding colleting dates. Dolejš (2015) was the first author to provide a collecting date for the specimens, the year 1930.

Regarding a standardized label. As a curatorial recommendation, an additional standardized label is proposed for all specimens, which should be labeled as syntypes following recommendation 72D ( ICZN, 1999). Scolopendra aztecorum Verhoeff, 1934 . SYNTYPE. MEXICO: Baja California Sur: La Paz (24.15070° N, 110.29724° W) or Los Inocentes (23.77716° N, 110.65430° W), under decaying cactuses. Leg: Dr. Jaroslav Štorkán. 1930.

Regarding specimens’ size: Verhoeff (1934) twice mentioned that he examined 4 adults and 2 juveniles. The adults were described as 118–125 mm in length, with 30–33 antennal articles and margination incomplete on T7 but marked on TT8–21. The juveniles were described as 70–77 mm in length, with 27–28 antennal articles and margination marked on TT14–21. However, it seems that Verhoeff (1934) mistook the numbers. Our data (table I) shows that Verhoeff (1934) categories are not absolute (juvenile RMNH.CHIL.154 has an antennal article count corresponding to the adult category) and that 2 syntypes are adults and 3 are juveniles. This makes us think that there were originally 2 adults and 4 juveniles, but without the sixth syntype this cannot be assured.

Figures

Figure 1. Map showing the type locality of Scolopendra aztecorum syntypes. Mexico, Baja California Sur, 1) La Paz and 2) Los Inocentes.

Figure 2a-h. Scolopendra aztecorum, syntype NMP P6E-1303. a) Head, dorsal. b) Tergite 1. c) Forciples. d) Toothplates. e) Tergite 21. f) Prefemur of leg pair 21, dorsal. g) Posterior segments, ventral. h) Posterior segments, lateral, right leg 20 showing the two tarsal spurs (ts).