JHR 82: 347-359 (2021) ge, JOURNAL OF meet snnscan na doi: 10.3897/jhr.82.64130 RESEARCH ARTICLE ) Hymenopter a 4 https://jhr.pensoft.net The Inarasional Society of Hymenoperiss, RESEARCH New synonymies in Sphecodes Latreille, 1804 (Hymenoptera, Apoidea, Halictidae) species from Italy, with taxonomic notes on type material Yulia V. Astafurova', Maxim Yu. Proshchalykin? | Zoological Institute, Russian Academy of Sciences, Saint Petersburg 199034, Russia 2 Federal Scientific Center of the East Asia Terrestrial Biodiversity, Far East Branch of the Russian Academy of Sciences, Vladivostok 690022, Russia Corresponding author: Maxim Yu. Proshchalykin (proshchalikin@biosoil.ru) Academic editor: Jack Neff | Received 8 February 2021 | Accepted 22 February 2021 | Published 29 April 2021 http://zoobank.org/9161 D8BF-832A-4DB5-B200-A7CD3C6B4BB5 Citation: Astafurova YuV, Proshchalykin MYu (2021) New synonymies in Sphecodes Latreille, 1804 (Hymenoptera, Apoidea, Halictidae) species from Italy, with taxonomic notes on type material. Journal of Hymenoptera Research 82: 347-359. https://doi.org/10.3897/jhr.82.64130 Abstract Type specimens of the bee genus Sphecodes Latreille, 1804 (Hymenoptera: Halictidae), described by Vit- torio Nobile and Giuseppe Fabrizio Turissi from Italy and deposited in the Zoologische Staatssammlung, Miinchen, Germany, were studied. Detailed information on the primary types of the six nominal species and their illustrations are provided. Sphecodes banaszaki Nobile & Turrisi, 2004, syn. nov., S. marcellinoi Nobile & Turrisi, 2004, syn. nov., S. walteri Nobile & Turrisi, 2004, syn. nov., S. iosephi Nobile & Turrisi, 2004, syn. nov., and S. tomarchioi Nobile & Turrisi, 2004, syn. nov., are all synonymized with Sphecodes combai Nobile & Turrisi, 2004, which is rediagnosed. Also accepted synonymy of S. campadellii Nobile & Turrisi, 2004 = S. geoffrellus (Kirby, 1802), previously suggested by M. Schwarz’s and F. Gusenleitner’s (2012). Keywords Anthophila, Apiformes, cleptoparasites, synonymy, taxonomy Copyright Yu. V.Astafurova, M. Yu. Proshchalykin. 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. 348 Yu.V. Astafurova & M.Yu. Proshchalykin / Journal of Hymenoptera Research 82: 347-359 (2021) Introduction One of controversial problems in the taxonomy of European species of the genus Sphe- codes Latreille, 1804 (Hymenoptera: Halictidae) is the status of the species described by Italian researchers V. Nobile and G.F. Turrisi (2004) from Italy. They described seven new species (each from a single male), attributing them to the Sphecodes pinguiculus Pérez, 1903 species group. It was not surprising that the description of several new species from this difficult genus at once in within a fairly well-studied European fauna (four of them from the same type locality) attracted attention. A few years later their type specimens were examined by Austrian researchers M. Schwarz and FE. Gusenleitner (2012). As a result, they were all synonymized with two other widespread Palaearctic species: S. campadellii Nobile & Turrisi, 2004 with S. geoffrellus (Kirby, 1802), and the other six species, S. combai Nobile & Turrisi, 2004, S. banaszaki Nobile & Turrisi, 2004, S. marcellinoi Nobile & Turrisi, 2004, S. walteri Nobile & Turrisi, 2004, S. zose- phi Nobile & Turrisi, 2004, and S. tomarchioi Nobile & Turrisi, 2004 with S. margi- natus Hagens, 1882. Nevertheless, a year later, that taxonomic action was challenged by the authors of the Italian species (Nobile and Turrisi 2013), who restored their valid status. Jakub Straka (Prague, Czech Republic) also studied these types; he was convinced that only S. combai is a distinct, valid species from the S. marginatus species group (personal communication of J. Straka); however, he did not publish that in the review of the European Sphecodes, see Bogusch and Straka (2012: 2). Sphecodes combai occurs in Italy from Sicily all the way north to the border with Switzerland, and was also found in Greece (Bogusch and Straka 2014). Thus, there are several points of view on the taxonomic status of the Sphecodes species described by Nobile and Turrisi (2004), so we decided to readdress this mat- ter. We reexamined the holotype specimens of S. combai, S. banaszaki, S. marcellinoi, S. walteri, S. iosephi, and S. tomarchioi in Zoologische Staatssammlung, Miinchen, Germany (ZSM). The holotype of S. campadellii was stated in the original description as being deposited in the Istituto di Entomologia, Universita di Bologna, Italy (IUB); however, it has not been found either there (personal communication of G.F. Turrisi) or in ZSM (personal communication of O. Schmidt), although the later depository was mentioned by Schwarz and Gusenleitner (2012) and Nobile and Turrisi (2013). We agree with the opinion of J. Straka about the validity of S. combai and also ac- cept M. Schwarz’s and F. Gusenleitner’s point of view that the holotype specimens of the other five Italian species (except S. campadellii) are conspecific with it, hence the proposed new synonymies. Materials and methods ‘The ventral surface of some flagellomeres bear a distinctive patch of sensilla trichodea A (sensu Argent and Svensson 1982), which we refer to as ‘tyloids’, these sensilla are easily observable under a microscope. Abbreviations F and T are used for flagellomere Italian Sphecodes 349 and metasomal tergum, respectively. Density of integumental punctures is described using the following formula: puncture diameter (in um) / ratio of distance between punctures to average puncture diameter, e.g., 15—20 um / 0.5-1.5. Specimens were studied with a Leica M205A stereomicroscope and photographs were taken with a combination of stereomicroscope (Olympus SZX10) and digital camera (Olympus OM-D and Canon EOS70D). Final images are stacked compos- ites using Helicon Focus 6 software. All images were post-processed for contrast and brightness using Adobe Photoshop. Taxonomy Sphecodes combai Nobile & Turrisi, 2004 Fig. 1 Sphecodes combai Nobile & Turrisi, 2004: 119-120, &. Holotype: 3, Italy, Lazio, Al- bani Hills, Cecchina, Poggio Ameno, 280 m, 1.X.1990, M. Comba, on Hedera helix L. (ZSM), examined. Sphecodes banaszaki Nobile & Turrisi, 2004: 120-121, 4. Holotype: G, Italy, Lazio, Albani Hills, communal forest of Albano, 480 m, 23.V1I.1995, M. Comba (ZSM), examined, illustrated in Fig. 2. — Syn. nov. Sphecodes marcellinoi Nobile & Turrisi, 2004: 121-122, 3. Holotype: , Italy, Sicily, Mount Etna, Piano Vetore, 1700 m (Ragalna), 2.V.1999, S. Tomarchio (ZSM), examined, illustrated in Fig. 3. — Syn. nov. Sphecodes walteri Nobile & Turrisi, 2004: 122-123, 3. Holotype: GS, Italy, Sic- ily, Mount Etna south slope, near da Milia, 1500 m (Ragalna), 10. VIII.1999, S. Tomarchio (ZSM), examined, illustrated in Fig. 4. — Syn. nov. Sphecodes iosephi Nobile & Turrisi, 2004: 123-124, ae Holotype: S, Sicily, Mount Etna, south slope, near da Milia, 1400 m (Ragalna), 10.VHI.1999, S. Tomarchio (ZSM), examined, illustrated in Fig. 5. — Syn. nov. Sphecodes tomarchioi Nobile & Turrisi, 2004: 124-125, Italy, 4. Holotype: 3, Sic- ily, Mount Etna, south slope, near da Milia, 1400 m (Ragalna), 10.VIII.1999, S. Tomarchio (ZSM), examined, illustrated in Fig. 6. — Syn. nov. Diagnosis. Morphologically, S. combai is similar to members of the S. miniatus Hagens, 1882 species group, i.e. S. creticus Warncke, 1992, S. haladai Warncke, 1992, S. laro- chei Warncke, 1992, S. marginatus Hagens, 1882, S. miniatus Hagens, 1882, S. nomi- oidis Pesenko, 1979, S. schwarzi Astafurova & Proshchalykin, 2014, and S. sandykachis Astafurova & Proshchalykin, 2018 (see Astafurova et al. 2018). However, this species clearly differs from other species of this group by F2 with tyloids covering the entire ventral flagellar surface (versus glabrous F2 or F2 with tyloids covering only the basal part of F2, Fig. 7B, C). This character was also pointed out by Nobile and Turrisi (2013) as a clearly distinguishing one from S. marginatus. Another important feature mentioned by these Italian researchers is the gonostylar shape. Although being very 350 Yu.V. Astafurova & M.Yu. Proshchalykin / Journal of Hymenoptera Research 82: 347-359 (2021) Figure |. Sphecodes combai Nobile & Turrisi, 2004, holotype, male A head, frontal view B vertex, dorsal view C metasoma, dorsal view D T1, dorsal view E genitalia, dorsal view F labels. Scale bars: 0.5 mm. similar in these two species, in S. marginatus it is relatively smaller and close to the triangular membranous portion of the gonostylus (Fig. 7D), distinguishes, S. combai from its trapezoidal membranous portion is larger (Fig. 1E). Thus, its shape is most similar to that in S. nomioidis (Fig. 7E), although the position of the gonostylus or Italian Sphecodes 351 aa o f M } ve £5 Wola ah e to LAS. » Coll Lar se 4 fe TVA | push iat Ste 7p O82 Cree ake Coviaan wate de pei ff Ayes A et y i Fs fj rae cote ce “ i Sphecodas a in oF O01 Figure 2. Sphecodes banaszaki Nobile & Turrisi, 2004, holotype, male A head, frontal view B vertex, dor- sal view C metasoma, dorsal view D T1, dorsal view E genitalia, dorsal view F labels. Scale bars: 0.5 mm. deformation of the genitalia in some specimens can significantly complicate proper usage of this feature. In fact, we could not clearly differentiate between S. combai and S. nomioidis, except by F2 tyloids (F2 is entirely glabrous in S. nomioidis). Based on antennal morphology, Nobile and Turrisi (2004, 2012) referred S. combai (and the other six species described in 2004) to the S. pinguiculus Pérez, 1903 species group (i.e., to the same group with S. pinguiculus and S. intermedius Bliithgen, 1923). 352 Yu.V. Astafurova & M.Yu. Proshchalykin / Journal of Hymenoptera Research 82: 347-359 (2021) E ) COLLEZIONE ETNA og 1700 | 3759 ee ine : 23} NICOLOSI - CT TOMARCHIOS._ Browsers, : ie | : S| * PIANO VETORE : : Saku Sietka dot sor sy, Slides at CG “1 oz-v- 1999 ‘wcome olotypus Be oss ¥4 TOMARCHIO S.LEG. ™avellino! sp. n. F ae we Ps EF Be a Nobile & Turrisi det. 2004 oe Figure 3. Sphecodes marcellinoi Nobile & Turrisi, 2004, holotype, male A head, frontal view B vertex, dor- sal view C metasoma, dorsal view D T1, dorsal view E genitalia, dorsal view F labels. Scale bars: 0.5 mm. Although S. pinguiculus and S. intermedius also have a similar T2 with tyloids usually covering the entire ventral flagellar surface (sometimes T2 is entirely or partially gla- brous in S. intermedius), S. combai differs from these species by a shorter antenna with flagellomeres being 1.1—1.2 times as long as wide (versus 1.4—1.7, Fig. 7A), a shorter Italian Sphecodes 359 ‘COLLEZIONE ~ Sey : ETNA SUD M 1500 5160 Holotypus NICOLOSI - CT | TOMARCHIO S._ Gl °G.DA MILIA “10 - vIIr - 1999 3 TOMARCHIO S.LEG. ° Cc waiteri sp. n. ieee F . we Nobile & Turrisi det. 2004 ~akub Straka del 2011 Figure 4. Sphecodes walteri Nobile & Turrisi, 2004, holotype, male A head, frontal view B vertex, dorsal view C metasoma, dorsal view D T1, dorsal view E genitalia, dorsal view F labels. Scale bars: 0.5 mm. vertex (the distance from top of the head to the upper margin of the lateral ocellus, being about two lateral ocellar diameters as seen in dorsal view, versus 2.5—3), the face above the antennal sockets with sparser and shorter pubescence partially obscur- ing the integument (versus dense long pubescence entirely obscuring the integument, 354 Yu.V. Astafurova & M.Yu. Proshchalykin / Journal of Hymenoptera Research 82: 347-359 (2021) SICILY ETNA SUD M 1400 .NICOLOSI - CT >C.DA MILIA 10 - VIII - 1999 TOMARCHIO S.LEG. LY cane n. Nobite & Turrisi det. 2004 : ese F Figure 5. Sphecodes iosephi Nobile & Turrisi, 2004, holotype, male A head, frontal view B vertex, dorsal view C metasoma, dorsal view D T1, dorsal view E genitalia, dorsal view F labels. Scale bars: 0.5 mm. Fig. 7A), and a more sparsely and finely punctate T1 with punctures separated by 2—5 puncture diameters (versus at most 2). So, possible placement of S. combai to the S. pinguiculus species group is doubtful and would be premature without taking into account the female morphological features. Besides, morphologically it is more similar Italian Sphecodes 355 ' oon ' >% ste | SICILY >| ETNA SUD M 1400 NICOLOSI - CT *2C.DA MILIA 10 - VIII - 1999 ~ Nobile & Turrisi det. 2004 TOMARCHIO S.LEG. comieary *» — F a) ees Figure 6. Sphecodes tomarchioi Nobile & Turrisi, 2004, holotype, male A head, frontal view B vertex, dor- sal view C metasoma, dorsal view D T1, dorsal view E genitalia, dorsal view F labels. Scale bars: 0.5 mm. to S. marginatus and S. nomioidis, so most likely S. combai belongs to the same group with the latter two species. Female. The female of S. combai has not been described, but we examined two from Italy (1 2, Emilia Romanga, Comacchio, VII.2001, M. Richa, and 1 9, Sic- 356 Yu.V. Astafurova & M.Yu. Proshchalykin / Journal of Hymenoptera Research 82: 347-359 (2021) Se iee allie re , Sao ‘ 3 a ne Ss it? ’ ‘D) ; a4 x OS 4 yee E oo Ef} ee : of i ae 4 Les 5 ee . Pils ws Figure 7. Sphecodes pinguiculus Pérez, 1903 (A), S. marginatus Hagens, 1882 (B, D), and S. nomioidis Pesenko, 1979 (C, E), males A head, frontal view B, C antennae, lateral view D, E genitalia, dorsal view. Scale bars: 0.5 mm. ily, Naxos Strand, 17.V.1961, J. Gusenleitner), both in the personal collection of M. Schwarz, Ansfelden, Austria) determined by J. Straka (in 2012) as S. combai. These specimens are very similar to S. marginatus, but with a relatively denser punctate ocel- lo-ocular area. It is likely that these two female specimens belong to S. combai, thus providing an additional indication that this species is likely to belong to the informal S. miniatus species group. However, we refrain from describing the female sex of this species based on specimens not in the same series with conspecific males. In addition, if the assumption that S. combai belongs to the S. miniatus species group is correct, then the female of S. combai may have very weak morphological differences from the other Italian Sphecodes 357 Table |. Differences of Sphecodes combai from other species of the S. miniatus species group described by Nobile and Turissi (2004). Characters F2 | Ocello-ocular | Upper half of frons Coloration of terga Type locality Holotype specimens (below ocelli) S. combai (Fig. 1) 1.1 ]ca 10 — 1-3 10-15 pm / red-brown with lighter marginal zones Lasio, Albani confluent—1 Hills S. banaszaki (Fig.2) ] 1.2] 10-15 pm/ | 10-15 pm/ less 0.5 | 5-12 um } T1-T2 dark red-brown with reddish marginal | Lasio, Albani 0.5—2 zones, 13—T4 black, T5 reddish Hills S. marcellinoi (Fig. 3) | 1.2 10-15 pm cal0um/0.5-2 |ca ; ce dark red-brown with reddish T1-T2 marginal | Sicily, Mount / 1-2 Etna S. walteri (Fig. 4) 1.2}cal0pm/ 1-3} calOpm/0.5-1 |ca5 — dark brown with reddish T1-T3 marginal Sicily, Mount 2-6 zones Etna S. iosephi (Fig. 5) 1.2] 10-15 pm/ |} 10-15 pm/ less 0.5 | ca 5 pm/} dark red-brown with reddish T1-T3 marginal | Sicily, Mount confluent—1 2-6 zones Etna S. tomarchioi (Fig. 6) | 1.2|ca 10 pm/ 1-2} 10-15 pm/ less 0.5 | ca 5 pm / dark brown with lighter T1-T3 marginal Sicily, Mount 3-6 zones Etna members of this group. Therefore, description of the other sex of S. combai may also require additional genetic evidence, which is now unfortunately lacking for both sexes. Synonymy. Most species of Sphecodes exhibit a considerable intraspecific variation. While studying numerous specimens from various parts of the Old World we have observed variation in the body size, sculpture, coloration, and sometimes degree of development of the tyloids and relative length of the flagellomeres. Intraspecific versus interspecific variation in body size Sphecodes is an interesting topic that warrants a fur- ther, thorough investigation, including using molecular methods. Indicated differences between the seven nominal species described by Nobile and Turrisi (2004) were based on sculpture (of the vertex, frons, T1), coloration (of the mandibles, antennae, metasoma and legs) and the gonostylar shape. We have reexam- ined the holotypes of S. banaszaki, S. tomarchioi, S. iosephi, S. walteri, S. marcellinoi, S. combai and concluded that the minor morphological differences between them can be considered as intraspecific variation, as also pointed out by Schwarz and Gusen- leitner (2012). The main differences between the primary type specimens examined are outlined in Table 1. The holotype of S. marcellinoi has a sparser punctation on the upper half of the frons; tergal coloration of the holotype of S. combai is lighter and reddish; the holotype of S. banaszaki is larger (6 mm versus 4—5 mm) and with coarser and denser T1 punctation; the holotype of S. iosephi has a denser punctate ocello-ocular area and mandibles medially dark yellow (not entirely black as incorrectly indicated in the original description). The slight differences in the gonostylar shape, as illustrated by Nobile and Turrisi (2004), are the result of a deformation and different position of the gonostylus relative to the axis of genitalia, see also Schwarz and Gusenleitner (2012). On the other hand, such a rare feature as F2 with tyloids covering the entire ventral flagellar surface not only distinguishes the six nominal species of Sphecodes described by Nobile and Turrisi (2004) from Italy from the other Palaearctic species quite well, but also more likely (also based on their morphological similarity) indicates their con- specificity within S. combai. 358 Yu.V. Astafurova & M.Yu. Proshchalykin / Journal of Hymenoptera Research 82: 347-359 (2021) Sphecodes geoffrellus (Kirby, 1802) Melitta geoffrella Kirby, 1802: 45, 2 (type locality: England). Sphecodes affinis Hagens, 1882: 224, 9, a (type locality: Germany). Sphecodes fasciatus Hagens, 1882: 224, ¢ (type locality: Germany). Sphecodes rimalis Pérez, 1903: 221, 9 (type locality: France). Sphecodes impunctatus Meyer, 1922: 171, & (type locality: Russia). Sphecodes silvicola Tsuneki, 1983: 60, 9 (type locality: Japan). Sphecodes shirozui Tsuneki, 1983: 53, e (type locality: Japan). Sphecodes kitamius Tsuneki, 1983: 55, J (type locality: Japan). Sphecodes campadellii Nobile & Turrisi, 2004: 118-119, Fig. 1, &. Holotype: 3, Italy, Emila Romagna, Zocca (Modena), 8.VII.1962 (collection of the Institute of En- tomology “G. Grandi”, University of Bologna, Italy). Junior subjective synonym of S. geoffrellus (Kirby, 1802) according to Schwarz and Gusenleitner (2012: 74). Remark. We have not reexamined the holotype of S. campadellii because it has not been found in [UB/ZSM and probably is lost. However, we agree with its synonymy with S. geoffrellus as previously suggested by Schwarz and Gusenleitner (2012) based on type studies. Acknowledgements We are grateful to O. Schmidt (ZSM) for her kind assistance during our work in the ZSM, to S. Triapitsyn (Riverside, USA) for checking the English grammar, and also to J. Straka and G.F. Turrisi for their personal communications on the subject. The com- ments of subject editor (J. Neff) and two reviewers (J. Straka and P. Bogusch) helped to improve the quality of this paper. This investigation was supported by the Russian Foundation for Basic Research (grant numbers 19—04—00027 and 20-54—44014) and the state research project AAAA-—A19-—119020690101-6. References Argent L, Svensson B (1982) Flagellar sensilla on Sphecodes bees (Hymenoptera, Halictidae). Zoologica Scripta 11: 45-54. https://doi.org/10.1111/j.1463-6409.1982.tb00517.x Astafurova YuV, Proshchalykin MYu, Schwarz M (2018) New and little known bees of the genus Sphecodes Latreille, 1804 (Hymenoptera: Apoidea: Halictidae) from Central Asia. 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