Redescription of Andrena savignyi SPINOLA, 1838 (Hymenoptera, Anthophila, Andrenidae)

Linzer biol. Beitr. 56/2 487-526 Jänner 2025 Redescription of Andrena savignyi SPINOLA, 1838 (Hymenoptera, Anthophila, Andrenidae) Anselm KRATOCHWIL A b s t r a c t : The types of Andrena savignyi described by SPINOLA (1838) are deposited in the Museo Regionale di Scienze Naturali (MRSN) in Turin (Italy) (Fulvio Giachino, MRSN, per litt. 2021). Spinola reported that he acquired three Egyptian females from the insect tradesman Dr M. Waltl (Passau, Germany) (SPINOLA 1838) (Fulvio Giachino, MRSN, per litt. 2021). The type specimens were collected by the entomologist and traveller M. Fischer in Egypt. A review by WOOD (2023) revealed that only one specimen could be assigned to A. savignyi. WOOD (2023) designated the lectotype in 2022, but without analysing the lectotype or the species in more detail. This study will add historical aspects of the Spinola collection. The distribution area of A. savignyi extends from the Canary Islands to the eastern border of India, which encompasses about 10,000 km. As the taxonomic rank of the populations outside of the lectotype (Egypt) has not yet been clarified, only the populations from Egypt will be examined morphologically and morphometrically in this study. The basis for this analysis was 25♀♀ and 12♂♂ collected between 1900 and 1999. An important aspect of this study was the clarification of the variation of individual characteristics. The results show that an extraordinarily large variation of individual characteristics can be found within Egypt in different populations, but also within one population. This applies, for example, to the extent of the reddish colouration of the tergites, different structures of the propodeum or pubescence colour. A detailed description of the male Egyptian specimens will be presented for the first time. The analysis of the genitalia of A. savignyi and a comparison with those of A. inaquosa WOOD, 2021 show that this species can be regarded as synonymous with A. savignyi. A morphological and morphometric analysis of A. savignyi in populations from the Canary Islands, Tunisia, Egypt and northern India and a comparison with the results obtained here will be published later (KRATOCHWIL, in prep.). Key words: Suandrena, taxonomy, systematics, nomenclature, Egypt. Introduction and background The Genoese entomologist Marquis Massimiliano Spinola di Tassarolo (1780-1857) described the mining bee Andrena savignyi in 1838 based on three specimens he acquired from the insect tradesman Dr M. Waltl (Passau, Germany). The specimens were collected in Egypt by the entomologist and traveller M. Fischer (SPINOLA 1838). GUSENLEITNER & SCHWARZ (2002) assumed the Museo Regionale di Scienze Naturali (MRSN) in Turin (Italy) to be the location of deposition, which indeed is correct (Fulvio Giachino, MRSN, per litt. 2021). The very brief typus description by SPINOLA (1838) considered only a few 488 characteristic features, mainly the body’s colouration and pubescence. This study will present and discuss some historical aspects of the Spinola collection and the first description of A. savignyi by SPINOLA (1838). WOOD’S (2023) examination of the three females revealed that only one specimen can be assigned to A. savignyi. The other two females have to be attended to Melitta aegyptiaca (RADOSZKOWSKI, 1891). WOOD (2023) designated the lectotype of A. savignyi in 2022. However, a detailed description of A. savignyi is not available yet. Due to the mixture of two species and the small amount of treated morphological features, SPINOLA’S (1838) description did not lead to a comprehensive view. DYLEWSKA (1983) characterised 20 females of A. savignyi but also included specimens from outside Egypt. Unfortunately, DYLEWSKA (1983) did not specify where these 20 specimens came from. It can be assumed that only a part of the specimens she analysed originated from Egypt. GUSENLEITNER & SCHWARZ (2002) also used specimens from outside Egypt for the characterisation of A. savignyi. Only MOUSTAFA & IBRAHIM (1965) briefly described females from Egypt, the species’ locus classicus. The analysis in this study is based on numerous morphological and morphometric features. Emphasis is given to their variation. A description of the males of A. savignyi is only available from WALKER (1871) under the synonym A. munda. According to WARNCKE (1967) and GUSENLEITNER & SCHWARZ (2002), the specimens from WALKER (1871) originating from Egypt have all been lost. An inventory by Joseph Monks (BMNH) revealed that no more specimens of the Walker collection can be found in the BMNH. SAUNDERS’ (1904) description of the males refers to specimens from the Canary Islands, and the description by DYLEWSKA (1983) to a specimen from Tunisia. A detailed description of the males of the Egyptian populations is presented here for the first time. The distribution range of A. savignyi extends from the Canary Islands to East India (GUSENLEITNER & SCHWARZ 2002, SCHUH et al. 2010, ASCHER & PICKERING 2021, WOOD & MONTFARED 2022, KRATOCHWIL, in prep.). The analysis of the Egyptian populations carried out here is necessary to clarify the taxonomic status of populations outside Egypt. Many species descriptions exist, which were later regarded as synonyms of A. savignyi. A review of the synonyms is in preparation (KRATOCHWIL, in prep.). Previous revisions of other Andrena taxa (A. tiaretta group, A. wollastoni group, A. cyanomicans group) have shown that cryptic species are also included within these widely distributed aggregates (KRATOCHWIL 2015, 2020; see also PRAZ et al. 2022). It can be assumed that cryptic species occur within the very large distribution area of A. savignyi. This study will mainly address the following questions: - Which historical facts are known about the species description of Andrena savignyi by SPINOLA (1838)? - Which characterisations of the females of A. savignyi have been published yet, and how can the new morphological and morphometric data improve the knowledge about this characterisation? - By which features can the males of A. savignyi be characterised morphologically and morphometrically? - To what extent are females and males characterised by individual variation? 489 Material and methods 1. Materials examined In this study, a total of 25♀♀ and 12♂♂ were analysed (see Appendix: Specimens examined). The specimens are deposited in the following collections (with acronyms for each depository) and individually characterised by an identity code (ID-No): MRSN .................. Museo Regionale di Scienze Naturali, Turin (Italy) OLML .................. Biologiezentrum Linz, Oberösterreichisches Landesmuseum, LinzDornach (Austria) SMF ..................... Naturmuseum Senckenberg Frankfurt (Germany) ZMHB ................. Museum für Naturkunde der Humboldt-Universität, Berlin (Germany) ZMS ..................... Zoologische Staatssammlung, München (Germany). Fig. 1 shows the 8 localities to which the specimens can be assigned. Fig. 1: Localities of the studied specimens of Andrena savignyi (21♀♀, 11♂♂). In addition, five further specimens were examined that were only labelled with "Egypt" (4♀♀, 1♂); Assuan = Aswan. 490 2. Morphological and morphometric analyses The following morphological features were analysed: structure, colour and pubescence of head (clypeus, labrum, labral process, mandible, scapus, antennal base, antenna, flagellomeres, frons, paraocular area, fovea facialis, vertex, genal area), mesosoma (mesoscutum, scutellum, propodeum, femur, tibia, basitarsus, mediotarsi, mesepisternum, propodeal corbiculae, trochanteral and femoral flocculus, tibial scopa, wing, pterostigma, pterostigmoid margin, veins) and metasoma (tergites, sternites, pygidium, genitals). Morphometric parameters were also analysed using terms and definitions according to MICHENER (2007) and KRATOCHWIL (2015, 2020, 2021a). The following parameters were analysed for females (in alphabetical order): basitarsus 3 length (lateral view, central distance), basitarsus 3 width (lateral view, central distance); clypeus width (maximal width from left to right side), clypeus length (central length from base to apex), clypeus puncture distance (average distance between the punctures), clypeus puncture diameter, clypeus width (maximal width); distitarsus width (dorsal view), distitarsus length (lateral view, central distance); eye length (maximal length of the eye, area with ommatidia, without eye ring, frontal view), eye width (maximal width of the eye, lateral view); femur 3 length (lateral view, central distance), femur 3 width (lateral view, central distance); flagellomere 1 length (lateral view, central distance); flagellomere 2 length (lateral view, central distance); flagellomere 3 length (lateral view, central distance); head length (from top of vertex to lower margin of clypeus), head width (maximal width in frontal view); interocular lower distance (between the left and right eyes at the lowest point, area with ommatidia, without eye ring), interocular upper distance (between the left and right eyes at the uppermost point, area with ommatidia, without eye ring); labrum process length, labrum process width apically, labrum process width basally; mediotarsus 3 segment 1, segment 2, segment 3 (lateral view, central distance); mesoscutum puncture distance, mesoscutum puncture width, mesoscutum length (central length between the beginning and the end of the mesoscutum), mesoscutum width (including tegulae); metasoma length (between the beginning of the 1st tergite and the apex of the pygidium), metasoma width (maximal tergite width); ocelloccipital distance (between posterior ocellus and preoccipital ridge); ocellus diameter (central ocellus without ocellus ring); propodeum basal area length (dorsal view), pterostigma length (maximum length between proximal base of pterostigma and position where the vein R1 has its typical width); tibia 3 length (lateral view, central distance), tibia 3 width (lateral view, central distance); wing length (maximal length of the forewing from the wing base to the apex). All analyses were carried out with a modular stereomicroscope Wild M3Z, Heerbrugg, Switzerland with a 25x eyepiece (16.25x, 40x, 62.5x and 100x). The program EAZYDRAW (2020) was used for drawings. Detailed photo documentation is available for all specimens studied (phototube for Wild M3Z with Canon EOS 7D Mark II, photo stacking over 30100 shots per image with FOTO Stacker Version 1.6 (29) (BOLTNEV & KACHER 2017). Unless otherwise stated, the photos were taken by the author. 3. Abbreviations Fl1, Fl2, etc. = first, second, etc. flagellomere; T1, T2, etc. = first, second, etc. metasomal terga; S1, S2, etc. = first, second, etc. metasomal sterna. 491 Results and discussion Historical aspects of the Spinola collection and the description of Andrena savignyi by SPINOLA (1838) The Genoese entomologist Marquis Massimiliano Spinola di Tassarolo (1780-1857) was a famous, scientifically active entomologist. He was born on 10 July 1780 in Pézenas (France) and died on 12 November 1857 in Tassarolo (Italy). He described hundreds of species of Coleoptera, Hymenoptera and Hemiptera (GESTRO 1915). He personally collected on his estates in Spain and South America and was in intensive dialogue with numerous European entomologists. He invested a lot of money in the purchase of insects and acquired parts or entire collections from other famous entomologists of his time, such as the French hymenopterologist Lepeletier (Amédée Louis Michel Le Peletier, comte de Saint-Fargeau) or the hemipterologist Jean-Guillaume Audinet-Serville (PASSERIN D’ENTRÉVES 1980, ROSA & XU 2014). Thanks to the extensive correspondence that Spinola kept, it was possible to reconstruct the collection in the 1980s (CASOLARI & MORENO 1978, CASOLARI & CASOLARI 1980, PASSERIN D’ENTRÉVES 1983). It was organised according to a classification system of the time: one or more specimens of a species were pinned together, their labels removed from the specimens and a main label attached underneath (Fig. 2). Depending on the origin of the species, Spinola chose different colours: white (Europe including Northwest Asia, the Near and Middle East), yellow (Asia), blue (Africa), green (North, Central and South America) or pink (Australia, Oceania) (Fulvio Giachino, MRSN, per litt. 2021). Each main label shows the genus and species name, the author of the species, the collector and the localities. The original Hymenoptera collection consisted of 69 boxes. The collection was later stored in new boxes, retaining the original arrangement and labelling (CASOLARI & MORENO 1978, CASOLARI & CASOLARI 1980). SPINOLA (1838) reported that he acquired Hymenoptera from Egypt from the insect tradesman Dr M. Waltl from Passau (Germany). Specimens were collected in Egypt by the entomologist and traveller M. Fischer. This purchase involved a whole collection of Hymenoptera, the description of which was published in a 110-page publication in the "Annales de la Société Entomologique de France, volume 7" (SPINOLA 1838). Among these specimens was a new species, which Spinola named Andrena savignyi after the French zoologist Marie Jules César Lelorgne de Savigny (1779-1851). He deliberately chose this name because J. C. Savigny belonged to a circle of 167 scientists, engineers and artists who accompanied Napoleon Bonaparte’s army to Egypt as part of a "Commission des sciences et des arts" during the "Egyptian campaign" (also known as the "Egyptian expedition") in 1798-1801. Although this "Egyptian expedition" largely failed militarily, it was a great success for natural science and for the research of the ancient Egyptian culture. The results of the expedition were published under the title "Description de l’Égypte" (1st edition 23 volumes, 2nd edition 37 volumes). J. C. Savigny and the French entomologist Victor Audouin are the authors of this expedition’s entomological results (JOMARD 1809). The collected Hymenoptera Anthophila were depicted true to scale and very accurately on copper engraving plates, whereby certain details (head, antenna, mouthparts, terminal tergite, tarsi, etc.) were also depicted in detail. Seven printed plates with bees include 492 Fig. 2: Box no. 138 of the Spinola collection. The specimens assigned by Spinola to Andrena savignyi are outlined in red. Only the specimen on the left side is A. savignyi. This specimen corresponds to the lectotype designated by WOOD (2023). The other two specimens are females of Melitta aegyptiaca (RADOSZKOWSKI, 1891). Photo: Fulvio Giachino, MRSN. 493 representatives of the genera Andrena, Anthophora, Ceratina, Colletes, Dasypoda, Eucera, Halictus, Hylaeus, Megachile, Nomada, Nomia, Osmia, Megachile and Sphecodes. However, a more precise taxonomic characterisation was not carried out prior to this publication. "Mr. Savigny presented a large number of species whose identification is really impossible without notes and coloured drawings. We had to confine ourselves to distinguishing the species on the basis of numbers..." (cited from JOMARD 1809). SPINOLA (1838) refers in his description to J. C. Savigny in JOMARD (1809) and to plate 7, no. 19, which in his opinion represents A. savignyi (Fig. 3) (MOUSTAFA 1986). Fig. 3: Title page of "Description de l’Égypte" (JOMARD 1809) and illustration of an Andrena bee, which, according to SPINOLA (1838), most probably represents the A. savignyi he described (JOMARD 1809). The lighter colour of the first three tergites in the illustration corresponds to the first three reddish-coloured tergites of A. savignyi. Picture credits: https://commons.wikimedia.org/wiki/ Category:Description_de_l%27Egypte._Histoire_naturelle._Tome_deuxième?uselang=de. The lectotype of Andrena savignyi in the Spinola collection of Turin GUSENLEITNER & SCHWARZ (2002) assume the Museo Regionale di Scienze Naturali (MRSN) in Turin (Italy) to be the location of deposition. Indeed, the type of Andrena savignyi is in the MRSN in Turin (Italy) (Fulvio Giachino, MRSN, per litt. 2021). Fig. 2 shows box no. 138 with the lectotype of A. savignyi.1 1 Box 138 contains some species that are not mentioned in GUSENLEITNER & SCHWARZ (2002), as well as other specimens undetermined by Spinola. In addition, there are other species described by Spinola that are now considered as synonyms: Andrena lanuginosa SPINOLA, 1837 as a synonym of A. (Campylogaster) pruinosa ERICHSON, 1835; A. rutila SPINOLA, 1838 (lectotype, WOOD 2023), and A. ephippium SPINOLA, 1838 (lectotype, WOOD 2023), as synonyms of A. (Melanapis) fuscosa ERICHSON, 1835 (WOOD 2023). Box 138 also contains species from the collection of Lepeletier, which Spinola acquired by purchase, e.g., A. lucida LEPELETIER, 1841 (today a synonym of A. limata SMITH, 1853). According to the checklist of GUSENLEITNER & SCHWARZ (2002) the museum of the type of A. lucida is assumed to be the Muséum National d’Histoire Naturelle, Paris (MNHN). However, it is not listed in the database there. There are four specimens of A. lucida, which most probably belong to the type series, three specimens from Oran (Africa) 494 Figs 4a-c show the three specimens described by Spinola as types and the label (outlined in red in the box). Spinola did not mark any types with a special label. As the specimens originate from Africa, the label should be blue, but it has become greenish over time (information, Fulvio Giachino, MRSN, 2021). The name chosen by Spinola is written on the label: Andrena Savignyi. Behind it is the letter "m", which stands for "mihi" (Spinola as author) and the female sign. Below this is the indication of the publication organ "Ann. soc. ent." (Annales de la Société entomologique de France, Paris). "D. Waltl" describes the origin of the specimens. "D." stands for "Dono" (Italian for "gift") or "Donavit" (Latin), and "Waltl" is the seller. Many specimens in the Spinola collection are labelled with such a "D" (information, Fulvio Giachino, MRSN, 2021). The label also describes the place of origin ("Égypte"). Further details can be found in the original letters of Spinola. The lectotype is dark-coloured on T3 (Fig. 4a); the first two tergites are reddish. Figs 4a-c: Lectotype of Andrena savignyi and the two Melitta aegyptiaca females; label of Spinola: (a) lectotype; (b, c) Melitta aegyptiaca; (d) label. For further explanations, see text (photos a, b, c, d: Fulvio Giachino, MRSN). Description of the females of Andrena savignyi by SPINOLA (1838) and descriptions by other authors Spinola published in 1838 in the "Annales de la Société Entomologique de France" (volume 7, page 512) the type description of A. savignyi. He did not mention that he had and one specimen from France. They all originate from Lepeletier and probably belong to the type series of this author (Fulvio Giachino, MRSN, per litt. 2021). 495 three specimens (females). SPINOLA (1838) characterised the female of A. savignyi as follows (translated from French): "Length 11.3 mm; width 3.4 mm. Antennae, body and legs black. Head, thorax and first abdominal segment covered with long white hairs. Three snow-white hair bands on the second, third and fourth segments. Denser pubescence on the fifth segment, white laterally and black in the centre. Hair of upper anal plate black; abdominal and leg hairs, including tibial brush, white; tarsi with golden yellow hairs. Wings hyaline; wing veins black. The first three segments of the abdomen are more or less reddish, but this colouration is variable. In some, three segments are predominantly red, in others black covers the red more or less, and occupies most of the second and third segments. Hairs on the seventh tergite black; hairs on the hind legs white." SPINOLA’S (1838) description is not very differentiated, as was usual for this time. It is based on a mixture of characteristics of A. savignyi and Melitta aegyptiaca. The abdominal hair bands of A. savignyi are not snow-white but rather yellowish-white, and the scopa is not white but golden. Melitta species resemble Andrena species, but among other features, their scopae are restricted to the posterior tibia and basitarsus, and their apical tarsi are broadened (Figs 4b, c). A. bipartita BRULLÉ (1839) was described based on specimens from the Canary Islands and renamed A. bicolorata SMITH, 1853 (nec Apis bicolorata ROSSI, 1790) due to the homonymy with A. bipartita LEPELETIER, 1841. COCKERELL (1938) also synonymised A. antilope PÉREZ (1895) from Algeria with A. bipartita. It is incomprehensible why COCKERELL (1938) did not use the valid name A. savignyi for the Egyptian populations. COCKERELL (1938) described Andrena bipartita aswanica from Egypt on the basis of a female collected by Capt. K. J. Hayward in Aswan (= Assuan, Egypt) on 29 January 1921 (Fig. 5b). In contrast to A. bipartita, COCKERELL (1938) characterised A. b. aswanica as follows: "Wings distinctly brownish, but not dark; second cubital cell somewhat contracted above, the hair, which is white in A. bipartita, is distinctly yellowish, first three tergites red." In an identification key, COCKERELL (1938) distinguished the nominate subspecies with the feature "2 tergites red", without mentioning what the nominate subspecies is and where it occurs. Fig. 5a shows a female from the Aswan region, collected by Mohamed A. Moustafa on 5 March 1972, and the type of A. bipartita aswanica (Fig. 5b). According to WARNCKE (1967), A. b. aswanica COCKERELL is synonymous with A. savignyi. DYLEWSKA (1983) listed the following characteristic features for A. savignyi in her description: body length 12-13.8 mm; index fovea facialis width to half facial width measured at the centre of the central ocellus = 0.33-0.36; clypeus length to clypeus width = 0.84; scapus length equal to the combined length of Fl1 and Fl2; Fl1 about 2.2 times longer than wide at the end; Fl2 and other flagellomeres about as long as wide, the last Fl about 1.5 times longer than wide; length-width ratio of maxillary palpal segments 1.6 : 2.8 : 1.8 : 1. 7 : 1.7 : 1.8, of the labial palpal segments 3.0 : 1.5 : 1.6 : 1.7; galea about 0.75 of the length of the clypeus; malar field about 2.6-2.7 times wider than the smallest length; genal field 1.9 times wider than the ocular profile; distance of the lateral ocelli from the posterior margin of the head about 1.6 ocelli size; labral process triangular and shiny; clypeus microsculptured, only at the base strongly shiny and coarsely punctured, distance between the punctures about 1-3 puncture diameters, with unpunctured line in the centre; mesoscutum finely microsculptured, strongly shining, only at the base with strong microsculpture and coarsely punctured; scutellum only microsculptured laterally and coarsely 502 The diameters of the punctures are 20-40 μm; the distances between them are 14-40 μm. There are several smaller, more closely spaced punctures at the base of the clypeus. The clypeus is characterised by a puncture-free midline in 68% (Figs 8a-d); in 32%, it is not puncture-free. Apically, the midline tapers into a triangular puncture-free area in more than half of all cases (Fig. 8b). Labral process:D YLEWSKA (1983) stated that the labral process is triangular and has a shiny surface. In the specimens studied, the labral process is not triangular, but always trapezoidal (Figs 9a-d). The measurements of the basal and apical width of the labral process confirm the trapezoidal shape: apical 0.17 mm (SD 0.02 mm, max. 0.21 mm, min. 0.14 mm), basal 0.70 mm (SD 0.02 mm, max. 0.74 mm, min. 0.66 mm). The length of the labral process is 0.21 mm (SD 0.01 mm, max. 0.24 mm, min. 0.19 mm). In 58%, the process is slightly apically rimmed, and in 38% clearly rimmed (Figs 9a, c). Only in 4% is it straight. In most cases, a shine is characteristic, and long rugulae (carinae) are visible (Fig. 9c). Differences in the shape of the labrum process also occur within a population. M a n d i b l e : The mandible is predominantly black in colour, often with a red to reddish-brown tip and a central area that may also be reddish (Fig. 9d). Figs 9a-d: Labral process: (a) emarginated (ÄG11; Luxor); (b) straight (ÄG 33; Aswan); (c) shiny with rugulae or carinae (ÄG8; Luxor); (d) mandible coloured red in the centre and apically (ÄG3; Fayed). 4.3 Mesosoma M e s o s o m a w i d t h a n d l e n g t h : The mesosoma, including the tegulae, has a width of 3.75 mm (SD 0.15 mm, max. 4.16 mm, min. 3.45 mm) and a length of 2.17 mm (SD 0.13 mm, max. 2.41 mm, min. 1.82 mm). Colour, sculpturing and puncturing:The colour of the mesosoma is black, as also indicated by SPINOLA (1838). As already mentioned, DYLEWSKA (1983) 503 considered specimens from outside Egypt in her characterisation, so these data are not comparable. The author characterises the mesoscutum as finely microsculptured, very shiny, with strong microsculpture only at the base, and coarsely punctured; the scutellum as microsculptured only laterally and coarsely punctured with a distance between the punctures of about 0.5-4 puncture diameters; and the postscutellum as coarsely microsculptured with a greasy shine and coarsely punctured, with a distance between the punctures of about 0.5-2 puncture diameters. However, a detailed analysis shows that the anterior mesoscutum is slightly shiny, shagreened, and densely and shallowly punctured (Fig. 10). The puncture diameter is 20 μm anteriorly, with a puncture distance of 27 μm. The mesoscutum is shiny from the centre to the end, not as densely but more deeply punctured (Fig. 10). The puncture diameter in the anterior area is 20 μm, with a puncture distance of 27 μm; in the remaining area, the puncture diameters are 14-26 μm, with puncture distances of 2741 μm. The scutellum is very shiny; the punctures have diameters of 14-41 μm and puncture distances of 14-27 μm (Fig. 10). Fig. 10: Mesoscutum weakly shiny anteriorly, shagreened, densely and shallowly punctured, shiny in the centre towards the end, not densely but more deeply punctured. The scutellum is strongly shiny, deeply punctured and with a larger distance between punctures (ÄG5; Egypt, without further localisation). P r o p o d e u m : The description of the propodeum in DYLEWSKA (1983) (roughly rugose, deeply fielded) and in GUSENLEITNER & SCHWARZ (2002) (fielded) is imprecise. The propodeum has a length of 0.58 mm (SD 0.03 mm, max. 0.65 mm, min. 0.54 mm). The structures are very variable (rugose, ridge-like, honeycomb-like, with longitudinal laminae and fan-shaped structures). There are transitions in the basal area from clear honeycomb-like structures and fielded structures to a dissolution of the structure to ridgelike structures (Figs 11a-d). Apically, ridge-like structures can also dominate, which can transition apically into a clear fan-shaped structure with laminae (Figs 11a, d) but also dissolve in structure. 504 There are four types, which occur in different frequencies: in 44%, there are some honeycomb-like structures basally, ridge-like structures or laminae towards the centre and a fanshaped structure apically. Basal honeycomb-like structures are absent in 36% of cases, but there are numerous ridge-like elements that form no or only a fragmentary fan-shaped structure apically. In 12% of cases, there are also no clear basal honeycomb-like structures, but there are ridge-like structures that also have a fan-shaped structure apically. Basal fragmentary honeycomb-like structures, otherwise only ridge-like structures and a very fragmentary fan-shaped structure, are found in 8% of cases. Figs 11a-d: Propodeum structures: (a) basal honeycomb-like pattern and apical fan-shaped structure (ÄG10; Egypt without further localisation); (b) basal honeycomb-like pattern and disintegrating fanshaped structure (ÄG4; Fayed); (c) no basal honeycomb-like pattern and missing apical fan-shaped structure (ÄG24; Luxor); (d) no basal honeycomb-like pattern and no distinct apical fan-sha ped structure (ÄG5; Egypt without further localisation). Colour of extremities:The information given by SPINOLA (1838) on the colouration of the extremities is not correct (confusion with Melitta aegyptiaca ). Femur 1 and tibia 1 are red-brown, tibia 1 apically orange-red. Basitar sus 1, mediotarsi and distitarsus are reddish-brown to orange-red; all tarsi are orange-red. Femur 3, tibia 3 and all tarsi of the hind extremities are red-orange (Figs 5a, b, Figs 6a, b). The m easurements of the posterior pair of legs yielded the values shown in Table 1. W i n g s , p t e r o s t i g m a a n d w i n g v e i n s : The wing length is 8.62 mm (S D 0.25 mm, max. 9.12 mm, min. 8.16 mm). SPINOLA (1838) characterises the wings as hyaline and the wing veins as black in colour. The wings of the examine d females are light translucent and not dark (Figs 12a-c). Only in very few cases is a very slight tint recognisable. The pterostigma shows a slight colour variation. In mo st cases, the pterostigma is yellow-orange to orange in the centre (88%) and yellow in a sma ller percentage of cases 505 (12%). T he margin of the pterostigma is yellow-reddish in 84%, reddish in 12%, and yellowish in 4%. The statement by MOUSTAFA & IBRAHIM (1965) that the pterostigma is coloured brown in the centre is not correct. The wing veins are in 84% yellow-reddish, in 12% yellow-orange and in 4% reddish in colour and in most cases characterised by dark margins. The information from SPINOLA (1838) that the wing veins are black is based on a confusion with Melitta aegyptiaca. Table 1: Lengths and widths of the hind extremities (femur; tibia; basitarsus; mediotarsus 1, 2 and 3; distitarsus) with standard deviations (SD), maximum and minimum values in mm. Pubescence:D YLEWSKA (1983) characterises the pubescence very undifferentiated (creamy white hairy dorsal thorax, legs and body). The information of SPINOLA (1838) does not correspond to the characteristics of A. savignyi (thorax covered with long white hairs; white abdominal and leg hairs, including the tibial scopa; tarsi with golden yellow hairs; white hairs on the hind legs) due to the confusion with the two specimens of Melitta aegyptiaca. The mesoscutum (Figs 5, 6, 10) is in most cases (68%) only marginally hairy. There are often only a few hairs in the centre. Seventy-six per cent have yellowish-white hairs, and 24% have whitish hairs. If the centre of the mesoscutum is hairy (32% of the specimens examined), 75% of the hairs are yellowish-white and 25% are white. The scutellum and postscutellum are hairier than the mesoscutum. In 84%, the pubescence is yellowish-white, in only 12% whitish, and in 4% yellowish-golden. Concerning the mesepisternum, 44% of the specimens had whitish and 36% yellowish-white pubescence. Twelve per cent showed pure white hairs; 8% had yellowish-white hairs in the upper part of the mesepisternum and white hairs in the lower part. The trochanteral flocculus and the femoral flocculus are identical in the percentage of hair colouration (whitish-yellow 88%; whitish 8%; white 4%). The tibial scopa is golden with only a few exceptions (92%; whitishyellow 8%). MOUSTAFA & IBRAHIM (1965) also refer to the golden hairs of the basitarsus. 506 Figs 12a-c: Wings, pterostigma and veins: (a) Wing hyaline, pterostigma in the centre reddish-orange with dark red margin, veins reddish with dark red margin (ÄG10, Egypt without further localisation); (b) wing slightly dark, pterostigma yellow-orange in the centre, becoming dark orange towards the margin with reddish margin, veins yellowish-orange with dark margin (ÄG16, Luxor); (c) wing hyaline, pterostigma yellow-orange in the centre with dark margin, veins yellow-orange with dark red margin (ÄG2, Wadi Digla). 507 4.4 Metasoma M e t a s o m a w i d t h a n d l e n g t h : The metasoma has a width of 3.68 mm (SD 0.18 mm, max. 4.00 mm, min. 3.20 mm) and a length of 6.29 mm (SD 0.43 mm, max. 7.04 mm, min. 5.36 mm). C o l o u r o f t e r g i t e s : The information on the tergite colouration of A. savignyi varies widely in the literature. According to SPINOLA (1838), T1-T3 are more or less reddish. The lectotype, however, has only two reddish tergites (Fig. 4a). DYLEWSKA (1983) emphasised that T1-T3 is completely or partially reddish in colour, which is confirmed by the results obtained here. GUSENLEITNER & SCHWARZ (2002) pointed out the orange-redcoloured T1 and T2 as well as occasionally equally coloured T3. C OCKERELL (1938) described A. bipartia aswanica from Egypt with three red tergites. MOUSTAFA & IBRAHIM (1965) characterised the abdomen only with basal orange colour. Tergite-specific information on colouration is not given. The statement by DYLEWSKA (1983) for the apical tergite depressions that they are translucent reddish is not quite correct, as they are clearly metallic greenish and orange. According to the results presented here, the tergites’ colouration and pattern show considerable variation (Figs 13, 14). The sequence of the examples follows an increasing red colouration: T1: Dorsally in 60% of the specimens, the colour is orange-reddish but not spotted (Figs 13e, f-h; Figs 14a, c, d, h), in 16% orange-reddish and slightly spotted (Figs 14b, c, e), in 16% orange-reddish but clearly spotted (Figs 13a-d, g; Figs 14b, e, g), and in 8% yellow-orange (Fig. 14f). Some specimens have a narrow brown-red margin apically on T1 in front of the depression (Figs 13b, e, f). On the front, 60% have a small area of reddish-brown colouring (Figs 13b, e-h; Figs 14b-d), 16% have a larger area of reddish-brown laterally, 16% are orange-reddish, similar to the centre (Fig. 13a, Fig. 14a), and 8% are yellow-orange (Fig. 15c). The punctures on dorsal T1 have a diameter of 20 μm and a puncture distance of 20 μm. The tergite surface is very shiny. T2: In 32%, T2 is orange-reddish and slightly or more strongly spotted (Figs 13a, c, e; Figs 14b, e); in 28%, it is only orange-reddish (Fig. 13f; Figs 14c, h); in 24%, it is orangereddish with a central more or less distinct spot (Figs 13d, g, h; Figs 14a, d, g); in 8%, orange-reddish with two central spots (Fig. 3b; Fig. 14e); and in 8%, uniformly yelloworange (Fig. 14f). Punctation and shine are as in T1. The depression is reddish-greenish iridescent. In 96%, there is a small black spot laterally (Figs 15a-c), and only 4% are without a spot. T3: Of all tergites, T3 shows the greatest variability in colouration and patterns. In 28%, black colour dominates, which is characterised by a bluish shine (Figs 13a, b, d; Fig. 14a) or, in some cases, reddish parts (Figs 13b, c), which are often restricted laterally. Twenty-eight per cent of T3s are orange-reddish (Fig. 14e), either completely or with a dark area in the centre (Figs 14a, d, g). Dark reddish-brown colouring dominates up to 20% (Figs 13g, h), wherein the centre may also have a dark area. Red to brown-reddish colouring dominates in 16% (Figs 13e, f; Fig. 14b), and yellow-orange in only 8% (Figs 14f, h). The base is coloured black with a bluish shine in 64% of cases (Figs 13b, e, g; Figs 14b, g), dark reddish-brown in 20% (Figs 13f, e), and orange-reddish to yellow-orange in 16% (Figs 14a, c, d). The apical margin is in 50% orange-reddish (Figs 13a, c-e, h; Figs 14a-e, g), in 29% reddish (Figs 13b, f, g), in 13% orange-reddish and metallic slightly greenish, and in 8% yellow-orange. The depression is s orange-greenish iridescent. The punctation corresponds to T2. 508 T4: In 56%, T4 is black with a bluish shine (Figs 13a, c-h; Fig. 14b); in 20%, black with a bluish shine but a red apical margin (Figs 14a, c, d); in 12%, also black with a bluish shine but laterally reddish brown and with a red apical margin (Figs 13b, g); in 12%, yelloworange to orange-reddish and apically somewhat reddish-brown (Figs 14e, f, h). The punctation corresponds to T2. The depression is uniformly metallic and greenish. T5: In 48%, T5 is black and reddish-brown (Figs 13a, b, d-h; Fig. 14a); in 24%, reddish-brown (Figs 14c, f, g); in 12%, bluish-black, and laterally reddish-brown (Fig. 13c; Figs 14b, d); in 12%, the base is black and metallic green with reddish depression (Fig. 14e); in 4%, yellowish-orange mixed with some reddish-brown areas (Fig. 14h). A total of 7 types of tergite colouration can be distinguished: -T1, T2: orange-reddish; T1, T2 partly spotted; T3, T4 black with bluish shine; T5 black to reddish-brown (Figs 13a, b). -T2: orange-reddish margin in front of the depression (Figs 13c, d). -T2, T3: reddish to brown-reddish, partly with a reddish margin (Figs 13e-h). -T2: orange-reddish, partly spotted (Figs 14a, b). -T3: orange-reddish, partly spotted (Figs 14c, d). -T4: orange-reddish (Figs 14e-g). -T5: yellow-orange with partial red-brown colouring (Fig. 14h). 509 Opposite: Figs 13a-h: Colouring and pattern of the tergites. Increase in the red-coloured portions of the tergites. (a) (ÄG1, Fayed): T1 orange-reddish with dark mottling, laterally like the centre; T2 orange-reddish, slightly mottled; T3 entire surface black with bluish shine, apically with narrow orange-reddish margin, depression orange-greenish iridescent; T4 black with bluish shine, depression metallic greenish; T5 black, reddish-brown; (b) (ÄG13, Luxor): T1 orange-reddish with a dark spot, small area reddishbrown anteriorly; T2 orange-reddish with two black spots; T3 basally bluish-black, centrally black, laterally dark reddish-brown, apically with narrow reddish margin, orange-greenish iridescent depression; T4 black with bluish shine, depression orange-greenish iridescent; T5 black and reddishbrown; (c) (ÄG6, Luxor): T1 orange-reddish, slightly mottled, small area reddish-brown anteriorly; T2 orange-reddish slightly mottled; T3 basally bluish-black, centrally black, reddish translucent, apically with narrow orange-reddish margin, depression orange-greenish; T4 black with bluish shine, depression metallic green; T5 bluish-black, laterally reddish-brown; (d) (ÄG2, Wadi Digla): T1 orange-reddish, slightly mottled, coloured anteriorly and laterally like centre; T2 orange-reddish with a black spot; T3 totally bluish-black, apically with narrow orange-reddish margin, depression orangegreenish iridescent; T4 black with bluish shine, depression metallic green; T5 black-reddish brown; (e) (ÄG8, Luxor): T1 orange-reddish, small area reddish-brown anteriorly; T2 orange-reddish, slightly mottled; T3 bluish-black basally, otherwise reddish or brown-reddish, apically with narrow orange-reddish margin, depression orange-greenish; T4 black with bluish shine, depression metallic green; T5 black-reddish-brown; (f) (ÄG9, Luxor): T1 orange-reddish, small area of reddish-brown colouring anteriorly and laterally; T2 orange-reddish; T3 basally reddish-brown, otherwise reddish or brown-reddish, apically with broad reddish margin, depression orange-greenish iridescent; T4 black with bluish shine, depression metallic green; T5 black-reddish brown; (g) (ÄG18, Luxor): T1 orange-reddish with central spot, small area reddish-brown anteriorly and laterally; T2 orangereddish; T3 basally bluish-black, centrally black, otherwise dark reddish-brown, apically with narrow reddish margin, depression orange-greenish iridescent; T4 black with bluish shine, depression metallic green; T5 black and reddish-brown; (h) (ÄG11, Luxor): T1 orange-reddish with central spot, small area reddish-brown anteriorly and laterally; T2 orange-reddish; T3 totally dark reddish-brown, apically with narrow orange-reddish margin, depression orange-greenish iridescent; T4 black with bluish shine, depression metallic green; T5 black and reddish-brown. Sculpturing and puncturing of the tergites: DYLEWSKA (1983) characterised the tergites as finely microsculptured, very shiny and with fine puncturing. The distance between the punctures is about 0.5-3.0 times the puncture diameter. According to our own measurements, the puncture diameter is usually 20 μm, with a puncture distance of 20 μm. Pubescence of tergites:S PINOLA (1838) pointed out that there are three snowwhite hair bands on T2-T4, with denser pubescence on T5 with white hairs laterally and black hairs in the centre. The hairs of T6 are described as black. Here, too, the description is based on a confusion with Melitta aegyptiaca. According to DYLEWSKA (1983), T1 has whitish hairs laterally, T2-4 broader whitish apical bands, and T5 and T6 brown-black hairs. A detailed analysis shows a large variation in the tergites’ hair colouration. - T1: yellowish-white patchy hairy (68%), whitish (16%), white (12%), patchy hairy (4%). - T2: basal yellowish-white patchy hairy (72%), whitish (16%), white (12%). - T3, T4: basal very short yellowish hairs (52%), basal very short whitish hairs (20%), whitish (16%), white (12%). - T5: central brown hairs, lateral yellow-white hairs (56%); central reddish-brown, lateral yellow-white hairs (20%); central reddish, lateral yellow-white hairs (16%); central brown, lateral white hairs (4%); central reddish, lateral white hairs (4%). - T6: reddish-brown hairs (64%), dark brown (24%), brown (12%). - T2-T4: white closed bands (64%), yellowish-white closed bands (36%). 510 Figs 14a-h: Colouring and pattern of the tergites. Increase in the red-coloured portions of the tergites. (a) (ÄG4, Fayed): T1 orange-reddish, anteriorly and laterally stained like centre; T2 orange-reddish with central dark reddish-brown area; T3 basally reddish and metallic slightly greenish, centrally black, apically with narrow orange-reddish margin, depression orange-greenish iridescent; T4 black with bluish shine, margin brown-red, depression metallic green; T5 black-reddish brown; (b) (ÄG7, Luxor): T1 orange-reddish, slightly mottled, small area reddish-brown anteriorly and laterally; T2 orangereddish, slightly mottled; T3 basally bluish-black, otherwise reddish to brown-reddish, apically with narrow orange-reddish margin, depression orange-greenish iridescent; T4 black with bluish shine, depression metallic green; T5 bluish-black, laterally reddish-brown; (c) (ÄG19, Aswan): T1 orangereddish, slightly reddish brown anteriorly; T2 orange-reddish; T3 basally and centrally orange-reddish, faint dark mottling, apically with narrow orange-reddish margin, depression orange-greenish iridescent; T4 black with bluish shine, laterally reddish-brown, marginally red, depression metallic green; T5 reddishbrown; (d) (ÄG5, Egypt): T1 orange-reddish, laterally slightly reddish brown anteriorly; T2 orangereddish; T3 basally and centrally orange-reddish, basally a small bluish-black spot, apically with narrow orange-reddish margin, depression dark greenish; T4 black with bluish shine, marginally red, depression metallic green; T5 bluish-black, laterally reddish-brown; (e) (ÄG3, Fayed): T1 orange-reddish, slightly mottled; T2 orange-reddish, slightly mottled; T3 basally two light reddish-brownish spots, otherwise orange-reddish, apically with narrow orange-reddish margin, depression dark greenish; T4 orangereddish, centre with reddish-brownish spot, depression metallic green; T5 base dark, depression metallic greenish; (f) (ÄG33, Aswan): T1 yellow-orange; T2 yellow-orange; T3 yellow-orange, depression orangegreenish iridescent; T4 yellow-orange, apically slightly reddish-brown, depression metallic green; T5 red-brown; (g) (ÄG22, Fayed): T1 orange-reddish, dark centrally, very slightly reddish brown laterally; T2 orange-reddish with a central spot; T3 base bluish-black, otherwise orange-reddish and dark centrally, margin orange-reddish and metallic greenish and orange, depression orange-greenish iridescent; T4 black with bluish shine, reddish-brown laterally and apically with red margin, depression metallic green; T5 reddish-brown; (h) (ÄG32, Egypt): T1 yellow-orange; T2 yellow-orange; T3 yelloworange, basally somewhat red-brown coloured, depression yellow-orange; T4 yellow-orange, apically somewhat red-brown, depression metallic green; T5 yellow-orange with somewhat red-brown colouration. 511 Figs 15a-c: Lateral view of abdomen: (a) (ÄG5, Egypt) T1 with small lateral reddish-brown colouration, T2 with a small lateral spot; (b) (ÄG12 Luxor) T1 with distinct lateral reddish-brown colouration, T2 with a large reddish-brown spot; (c) (ÄG3 Fayed) T1 with lateral yellowish-orange colouration, T2 with large reddish-brown spot. Figs 15d-e: Pygidium: (d) (ÄG4, Fayed); (e) (ÄG5, Egypt). Pygidium:According to GUSENLEITNER & SCHWARZ (2002), the pygidium is apically truncated, which is correct. The pygidium also varies in colour (Figs 15d, e). It is in 84% black, in 8% reddish-brown, in 4% black and reddish-brown, and in 4% black with a reddish tip. 518 Colour, sculpturing and puncturing:The colour of the mesosoma is black, slightly shiny in the anterior shagreened area, and densely and shallowly punctured. The mesoscutum is shiny from the centre to the end, not as densely but more deeply punctured. The puncture diameter is 27 μm in the anterior third, with a puncture distance of 13 μm; in the remaining area, the puncture diameter is also 27 μm, and the puncture distances are 27-60 μm. The scutellum is very shiny, deeply punctured in the centre with a distance of up to 67 μm, and more densely spaced towards the sides. P r o p o d e u m : The propodeum has a length of 1.35 mm (SD 0.07 mm, max. 1.46 mm, min. 1.24 mm). The structures are also very variable in males (ridge-like, with rugulae or laminae, honeycomb-like, and fan-shaped structures). There are transitions in the basal area from clear honeycomb structures to a dissolution of the honeycomb-like structure to ridge-like structures (Figs 20a-f). Apically, ridge-like structures can also dominate, which can transition into a clear fan-shaped structure (Figs 20a, b). There may also be a dissolution of the clear structures. Colour of extremities:G USENLEITNER & SCHWARZ (2002) only refer to the red colouration of the tarsi of the 3rd pair of legs. Femur 1 and tibia 1 are reddish-brown, as is basitarsus 1. The mediotarsi and the distitarsus are reddish-brown to reddish; the tarsi are reddish. The femur of the third pair of legs is reddish-brown; the tibia is reddish-brown to reddish. The tarsi are in 36% reddish-brown and in 64% orange-red (Fig. 17). The measurement of the posterior pair of legs yielded the values shown in Table 2. Table 2: Lengths and widths of the segments of the 3rd pair of legs (femur; tibia; basitarsus; mediotarsus 1, 2 and 3; distitarsus), with standard deviations (SD), maximum and minimum values in mm. 519 W i n g s , p t e r o s t i g m a a n d w i n g v e i n s : The wing length is 7.55 mm (SD 0.24 mm, max. 7.70 mm, min. 7.00 mm). The wings are light, translucent and not tinted (Fig. 17). The pterostigma shows a slight colour variation. In most cases, it is yellowish in the centre (64%), and orange in a smaller percentage of cases (36%). The margin of the pterostigma is always reddish. The wing veins are in 64% yellowish-orange and in 36% orange. Pubescence:According to W ALKER (1871), the thorax is white-haired. The specimens studied here show a variation in hair colouration: the mesoscutum, scutellum and postscutellum are in 64% yellowish-white and in 36% yellowish-golden, while the mesepisterum has whitish hairs in 64% and yellowish-white hairs in 36%. 6.4 Metasoma W i d t h a n d l e n g t h : The metasoma has a width of 3.29 mm (SD 0.20 mm, max. 3.36 mm, min. 2.96 mm) and a length of 5.75 mm (SD 0.44 mm, max. 6.16 mm, min. 5.20 mm). Colour of tergites:G USENLEITNER & SCHWARZ (2002) mentioned that in males, the red colouration of the tergites is strongly reduced and only the depressions are lightened. According to the results presented here, the colouration and patterns of the tergites also show considerable variation in the males (Fig. 21). This results in the following characterisation: -T1: In 45%, T1 is black to red-brown and has a red terminal margin. In 27%, the red colouring of the terminal margin is absent. In 18%, T1 is bluish-black to reddishbrown, sometimes with a red end margin. In 9%, T1 is reddish-brown with a red end margin. In all cases, the depressions of T1 are reddish-brown and metallic greenish. -T2: In 46%, T2 is black to reddish-brown and has reddish, usually metallic greenish areas laterally. Thirty-six per cent of specimens have a reddish end margin in addition to the black to reddish-brown colouring. In only 9% is T2 purely black to reddish brown, and in another 9%, it is bluish-black. In all cases, the depressions of T2 are reddish-brown and metallic greenish. -T3: In 64% of the specimens, T3 is black to reddish-brown, one-third of them with reddish areas laterally, one-third with a red end margin, and one-third only black to reddish-brown. Thirty-six per cent of specimens show a black colour with a bluish shine. In all cases, the depressions of T3 are reddish and metallic greenish. -T4: In most cases (46%), T4 is black with a bluish shine, but in 27% with a greenish shine. In 18%, T4 is black to reddish-brown, sometimes with a reddish lateral colour or with a red end marginally. Only in 9% is T4 completely bluish-black. The depressions of T4 are reddish and metallic greenish. -T5: Like T4, T5 is in 55% black with a bluish shine, but only in 18% with a greenishbluish shine. T5 is in 18% black to reddish-brown, either with reddish spots laterally or with a red end margin. In 9%, T5 is red to reddish-brown. The depressions of T5 are reddish and metallic greenish. Sculpturing and puncturing of the tergites:The puncture diameter is usually 20 μm, with a puncture distance of 20 μm. 520 Figs 21a-h: Colouration and pattern of the abdomen. Increase in the red-coloured portions of the tergites. (a) (ÄG27, Fayed): T1 bluish-black to reddish-brown, depression reddish-brown, metallic greenish; T2-T5 bluish-black, depressions reddish, metallic greenish; pygidium reddish; (b) (ÄG26, Cairo): T1 black to reddish-brown, red terminal margin, depression reddish-brown, metallic greenish; T2 as T1; T3 black, red terminal margin; T4, T5 black with red terminal margin, with bluish shine; T2-T5 depressions reddish, metallic greenish; pygidium reddish; (c) (ÄG30, Fayed): T1 black to reddish-brown, depression reddish-brown, metallic greenish; T2 black to reddish-brown, reddish spots laterally, depression reddish-brown, metallic greenish; T3-T5 black with bluish shine; T2-T5 depressions reddish, metallic greenish; pygidium reddish; (d) (ÄG35, Luxor): T1 black to reddishbrown, depression reddish-brown, metallic greenish; T2, T3 black to reddish-brown, laterally reddish and metallic greenish, depressions reddish-brown, metallic greenish; T4-T5 black with bluish shine; T2-T5 depressions reddish, metallic greenish; pygidium reddish; (e) (ÄG23, Luxor): T1 blackish to reddish brown, red terminal margin, depression reddish-brown, metallic greenish; T2 as T1, but T2 with broader red terminal margin; T3 blackish to reddish-brown; T4, T5 black with greenish-bluish shine; T2-T5 depressions reddish, metallic greenish; pygidium reddish; (f) (ÄG28, Fayed): T1 bluishblack to reddish-brown, partial red terminal margin, depression reddish-brown, metallic greenish; T2 black to reddish brown, depression reddish-brown, metallic greenish; T3-T5 black with bluish shine; T2-T5 depressions reddish, metallic greenish; pygidium reddish; (g) (ÄG20, Suez): T1 black to reddish-brown, red terminal margin, depression reddish-brown, metallic greenish; T2, T3 as T1, but T2 with broad red terminal margin; T4, T5 black with greenish-bluish shine; pygidium reddish; (h) (ÄG31, Fayed): T1 black to reddish-brown, red terminal margin, depression reddish-brown, metallic greenish; T2 black to reddish-brown, laterally reddish and metallic greenish, depression reddishbrown, metallic greenish; T3-T5 black with bluish shine, depressions reddish, metallic greenish; pygidium reddish. 521 P u b e s c e n c e o f t e r g i t e s : A detailed analysis shows the following variation of hair colouration of the tergites: -T1: yellowish-white with patchy hairs (68%), whitish (16%), white (12%), patchy hairs (4%). -T2: basally patchy hairs that are yellowish-white (72%), whitish (16%), white (12%). -T3, T4: basally very short yellowish hairs (52%), very short whitish hairs (20%), not short whitish hairs (16%), not short white hairs (12%). -T5: centrally brown, laterally yellow-white hairs (56%); centrally reddish-brown, laterally yellow-white hairs (20%); centrally reddish, laterally yellow-white hairs (16%); centrally brown, laterally white hairs (4%); centrally reddish, laterally white hairs (4%). -T6: reddish-brown (64%), dark brown (24%), brown hairs (12%). -T2-T4: white hair bands (64%), yellowish-white hair bands (36%). P y g i d i u m : In contrast to the females, the pygidium does not vary in colouration. It is reddish (Figs 15d, e). Figs 22a-f: Male genitalia: (a) ÄG29, Fayed; (b) ÄG 31, Fayed; (c) ÄG35, Luxor; (d) ÄG20, Suez; (e) ÄG21, Luxor; (f) ÄG 26, Cairo. 522 G e n i t a l i a : This is the first description of the male genitalia of A. savignyi from Egypt. Genitalia can also have a slight variation in colouration and shape (Figs 22a-f). The genitalia are relatively broad. The gonocoxes can be black, reddish-brown or reddish in colour. The relatively long apically arranged spines on the gonocoxes, which diverge slightly, are conspicuous. GUSENLEITNER & SCHWARZ (2002) characterised them as distinct but short and narrow. The gonostyli are indented, as is typical for the genus Suandrena, and have yellowishwhite to golden hairs on the apical margin (see also GUSENLEITNER & SCHWARZ 2002). The penis valve is extraordinarily broad and flat with lateral thickenings. In KRATOCHWIL (2021b) the genitalia of Andrena cyanomicans PÉREZ, 1895, A. fratella WARNCKE, 1968, A. maderensis COCKERELL, 1922, A. mirna WARNCKE, 1969, A. notata WARNCKE, 1968, and A. portosanctana COCKERELL, 1922 (all Suandrena species) are illustrated in Fig. 24. WOOD (2021) published photos of the genitalia of A. planiventris DOURS, 1872, and the newly described A. inaquosa from Palmyra (Iran). The morphological comparison shows that the genitalia of A. inaquosa are identical to those of A. savignyi. Thus, the name A. inaquosa is a synonym, which was already assumed in KRATOCHWIL (2021b). A more detailed morphological and morphometric analysis of the type at the OLML is planned. A male from the OLML, collected in Syria in 1899 (A. savignyi, det. Warncke), was analysed by A. Kratochwil. Synopsis There is only one specimen of Andrena savignyi which can be regarded as a lectotype. The previous descriptions of the female of A. savignyi are extremely incomplete and are not sufficient to describe the species with morphological precision. A description of the male of A. savignyi was not available at all yet. An analysis of a multitude of morphological and morphometric parameters and a sufficient number of females and males of different populations from the locus classicus (Egypt) was therefore necessary. The results of this study show that an extraordinarily large variation of individual characteristics can be found within Egypt in different populations, but also within one population. This applies, for example, to the extent of the red colouration of the tergites, different structures of the propodeum and the pubescence colour. Nevertheless, in addition to features that clearly characterise the subgenus Suandrena (WARNCKE, 1968), there are also many features that clearly distinguish A. savignyi from the other Suandrena species despite variation (see KRATOCHWIL 2021b). On the basis of these results, it is now possible to make comparisons with other populations of A. savignyi outside Egypt, within the overall distribution area from the Canary Islands to the western border of China. In particular, multivariate morphometric analyses will have to be used. These broad comparisons will be presented soon (KRATOCHWIL, in prep.). Acknowledgements I would like to thank Fulvio Giachino, MRSN, for taking photos of the lectotype of Andrena savignyi and other specimens from the Spinola collection. He also provided me with valuable information about Spinola and the collection. I am indebted to Joseph Monks (NHMUK), who checked the collection concerning specimens from Francis Walker. Fritz Gusenleitner, Esther Ockermüller and 523 Martin Schwarz (OLML); Stefan Schmidt (ZMS); Steffen Pauls and Jennifer Steppler (SMF); Michael Ohl and Stefanie Krause (ZMHB) lent me specimens from the named museums for the morphological and morphometrical analyses. I would like to thank them for their great co-operation. Zusammenfassung Das Typus-Exemplar der von SPINOLA (1838) beschriebenen Art Andrena savignyi ist im Museo Regionale di Scienze Naturali (MRSN) in Turin (Italien) hinterlegt (Fulvio Giachino, MRSN, per litt. 2021). Spinola berichtete, dass er drei ägyptische Weibchen von dem Insektenhändler Dr. M. Waltl (Passau, Deutschland) erwarb (SPINOLA 1838) (Fulvio Giachino, MRSN, per litt. 2021). Die Tiere wurden von dem Entomologen und Reisenden M. Fischer in Ägypten gesammelt. Eine Überprüfung durch WOOD (2023) ergab, dass nur ein Exemplar A. savignyi zugeordnet werden kann. WOOD (2023) benannte den Lectotypus im Jahr 2022, ohne jedoch diesen oder die Art näher zu analysieren. Im Rahmen dieser Untersuchung werden historische Aspekte der Spinola-Sammlung beschrieben. Das Verbreitungsgebiet von A. savignyi erstreckt sich von den Kanarischen Inseln bis zur Ostgrenze Indiens, was einer Strecke von etwa 10.000 km entspricht. Da der taxonomische Rang der Populationen außerhalb des Verbreitungsgebietes des Lectotyps (Ägypten) noch nicht geklärt ist, werden in dieser Studie nur die Populationen aus Ägypten morphologisch und morphometrisch untersucht. Die Grundlage für diese Analyse bildeten 25♀♀ und 12♂♂, die von 1900 bis 1999 gesammelt wurden. Ein wichtiger Aspekt war die Klärung der Variation der einzelnen Merkmale. Die Ergebnisse zeigen, dass eine außerordentlich große Variation individueller Merkmale auch innerhalb einer Population zu finden ist. 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Address of the author: Prof. Dr. Anselm KRATOCHWIL Department of Biology/Chemistry, Ecology Section University of Osnabrück Barbarastr. 13, D-49069 Osnabrück, Germany E-mail: [email protected] Appendix Specimens examined Details in the following order: Identification number (ÄG1, ÄG2……), collection (abbreviations see chapter "Material and Methods"), subcollection, sex, locality, date of detection, collector (leg.), determinator 1 (det.), determinator 2 (det.), used synonym, further comments. All specimens were checked by A. Kratochwil and as Andrena savignyi confirmed. ÄG1: OLML, coll. K. Warncke, ♀, Fayed?, leg. H. Priesner, det. Grünwaldt (A. savignyi); ÄG2: OLML, coll. K. Warncke, ♀, Wadi Digla, March, leg. Dr. H. Priesner, det. Warncke (A. savignyi); ÄG3: OLML, coll. K. Warncke, ♀, Fayed?, leg. H. Priesner, det. Warncke (A. savignyi); ÄG4: OLML, coll. K. Warncke, ♀, Fayed, February1943, leg. H. Priesner, det. Warncke (A. savignyi); ÄG5: OLML, coll. K. Warncke, ♀, leg. Vaino? Pekkola, det. H. Teunissen 1986 (A. savignyi); ÄG6: SMF, coll. A. Weis, ♀, Luxor, 17.03.1910, leg. Jacquet, det. Grünwaldt (A. savignyi), det. Moustafa (A. savignyi); ÄG7: SMF coll. A. Weis, ♀, Luxor, 17.03.1910, leg. Jacquet, det. Grünwaldt (A. savignyi), det. Moustafa (A. savignyi); ÄG8: SMF, coll. A. Weis, ♀, Luxor, 17.03.1910, leg. Jacquet, det. Grünwaldt (A. savignyi), det. Moustafa (A. savignyi); ÄG9: SMF, coll. A. Weis, ♀, Luxor, 17.03.1910, Jacquet, det. Grünwaldt (A. savignyi), det. Moustafa (A. savignyi); ÄG10: SMF, ♀, 06.03., without year, leg. A. Andres, det. Alfken 1924 (A. savignyi), det. Moustafa (A. savignyi), in the soil in the garden; ÄG11: SMF, coll. A. Weis, ♀, Luxor, 17.03.1910, leg. Jacquet, det. Grünwaldt (A. savignyi), det. Moustafa (A. savignyi); ÄG12: SMF, coll. A. Weis, ♀, Luxor, 17.03.1910, leg. Jacquet, det. Grünwaldt (A. savignyi), det. Moustafa (A. 526 savignyi); ÄG13: SMF, coll. A. Weis, ♀, Luxor, det. Schmiedeknecht (A. antilope), det. Moustafa, (A. savignyi); ÄG14: SMF, ♀, Wadi Riched, 01.03.1914, leg. A. Andres, det. Alfken 1925 (A. savignyi), det. Moustafa (A. savignyi); ÄG15: SMF, ♀, March, leg. A. Andres, det. Alfken 1925 (A. savignyi), det. Moustafa (A. savignyi), nesting in the soil in a garden; ÄG16: SMF, coll. A. Weis, ♀, Luxor, 1904, leg. Schmiedeknecht, det. Schmiedeknecht (A. antilope), det. Moustafa (A. savignyi); ÄG17: SMF, coll. A. Weis, ♀, Luxor, 23.03.1999, det. Friese 1900 (A. antilope); det. Moustafa; ÄG18: SMF, coll. A. Weis, ♀, Luxor, 17.03.1910, leg. Jacquet, det. Grünwaldt (A. savignyi), det. Moustafa (A. savignyi); ÄG19: SMF, coll. A. Weis, ♀, Assuan, 14.02.1910, leg. Jacquet, det. Grünwaldt (A. savignyi), det. Moustafa (A. savignyi); ÄG20: SMF, coll. A. Weis, ♂, Suez, Aegyptus 1890, 11.04.1896, Friese 1900 (A. antilope), det. Moustafa (A. savignyi); ÄG21: SMF, coll. A. Weis, ♂, Luxor, 17.03.1910, leg. Jacquet, det. Grünwaldt (A. savignyi), det. Moustafa (A. savignyi); ÄG22: ZMS, ♀, Fayed, February 1943, leg. H. Priesner; ÄG23: ZMS, coll. A. Weis, ♂, Luxor, 17.03.1910, leg. Jacquet, det. Grünwaldt (A. savignyi), det. Moustafa (A. savignyi). ÄG24: ZMS, coll. A. Weis, ♀, Luxor, 17.03.1910, leg. Jacquet, det. Grünwaldt (A. savignyi); ÄG25: ZMS, coll. A. Weis, ♀, Luxor, 17.03.1910, ♂, leg. Jacquet, det. Grünwaldt (A. savignyi); ÄG26: ZMS, ♂, Kairo, leg. Staudinger?; ÄG27: ZMS, ♂, Fayed?, leg. H. Priesner, det. Grünwaldt (A. savignyi); ÄG28: ZMS, ♂, Fayed, February1943, leg. H. Priesner, det. Grünwaldt (A. savignyi); ÄG29: ZMS, ♂, Fayed, February 1943, leg. H. Priesner, det. Grünwaldt (A. savignyi); ÄG30: ZMS, ♂, Fayed, leg. H. Priesner, Grünwaldt (A. savignyi); ÄG31: ZMS, ♂, Fayed, leg. H. Priesner, det. Grünwaldt (A. savignyi); ÄG32: ZMHB, ♀, leg. S. Ehrenberg, det. Moustafa (A. savignyi), Nr. 2546, old determination label ‘savignyi’; ÄG33: ZMHB, coll. Moustafa, ♀, Aswan District, 05.03.1972, leg. Moustafa, det. Moustafa (A. savignyi); ÄG34: OLML, coll. K. Warncke, ♀, det. Moustafa (A. savignyi), 2 old determination labels ‘A. antilope’, 1 old determination label ‘savignyi’; ÄG35: OLML, coll. K. Warncke, ♂, Luxor, det. Warncke; ÄG36: OLML, coll. K. Warncke, ♀, Sinai, Mitla Pass, 13.04.1973, leg. Bytinski-Salz, det. Warncke. ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Linzer biologische Beiträge Jahr/Year: 2025 Band/Volume: 0056_2 Autor(en)/Author(s): Kratochwil Anselm Artikel/Article: Redescription of Andrena savignyi SPINOLA, 1838 (Hymenoptera, Anthophila, Andrenidae) 487-526