Strange loves: a remarkable case of aberrant copulation in beetles (Coleoptera: Meloidae, Chrysomelidae)

A case of copulation between two mimic and repellent beetle species (a male of Timarcha fracassii, and a female of Meloe autumnalis), belonging to distinct families (Chrysomelidae, Meloidae), is recorded

The cranial apparatus glands of the canthariphilous Pyrochroa coccinea (Coleoptera: Pyrochroidae: Pyrochroinae), and their implications in sexual behaviour

Some Pyrochroidae species are known as "canthariphilous" for their attraction to cantharidin (CTD), a toxic terpene with anti-predatory effects, produced in nature by only two beetle families (Meloidae and Oedemeridae). It has been demonstrated that males of Neopyrochroa flabellata ingesting CTD are positively selected by females. Indeed, the compound is re-emitted from a glandular cranial apparatus as secretions that are licked up by females during courtship behaviour, inducing copulation. Herein, we provide the first description of the glands associated to the cranial apparatus of male Pyrochroinae using the European species Pyrochroa coccinea as a model. Morphological analyses show that the cranial apparatus consists of a concave pit lined with short setae retaining secretions emitted through numerous glandular pores. Ultrastructural investigations reveal the presence of two different class 3 glands (Gl.A and Gl.B), intermixed at the level of the pit but exhibiting distinct features. Gl.A are mainly characterised by short conducting canals, rounded nuclei and electrondense vesicles while Gl.B are characterised by long conducting canals, irregular nuclei, vesicles containing a particulate substance and a multifolded plasma membrane. Observations of sexual behaviour are also reported for P. coccinea and compared to N. flabellata, confirming the involvement of cranial apparatus secretions in courtship behaviour

Larval features of the Italian endemic Pyrochroa serraticornis kiesenwetteri Fairmaire, 1849 (Coleoptera: Pyrochroidae: Pyrochroinae) solve taxonomic uncertainties

Recent findings rearranged the taxonomy of the European species of the genus Pyrochroa Geoffroy (Coleoptera: Pyrochroidae), and to date three species are recognized: P. bifoveata Molfini et al., 2022 and P. coccinea (Linnaeus, 1761) with cryptic adults and distinctive larvae, and the polytypic P. serraticornis (Scopoli, 1763), including the subspecies kiesenwetteri Fairmaire, 1849. Incongruences between molecular and morphological analyses questioned the recognition of P. s. serraticornis and P. s. kiesenwetteri as taxa of the same species. In the present paper, observations of larval characters of P. s. kiesenwetteri confirm this taxon as a subspecies of P. serraticornis. Moreover, new characters of P. s. serraticornis larvae are offered by analysing specimens from a new European locality

Phylogeny of European Pyrochroa (Coleoptera, Pyrochroidae) reveals cryptic taxa and different glacial histories

Only three saproxylic species of Pyrochroinae (Coleoptera: Pyrochroidae) are distributed in Europe, two of which belonging to Pyrochroa: P. coccinea and P. serraticornis. However, P. serraticornis is polytypic, for the presence of the endemic subspecies P. s. kiesenwetteri in southern Italy. Using both molecular and morphological data, we explored the phylogeny of the European Pyrochroa species. A multilocus (COI, CAD, 28S) phylogenetic analysis helped highlight different evolutionary histories for the two examined species. First, P. coccinea, distributed throughout Europe, showed a high differentiation among Italian and European populations. Furthermore, three different taxonomic entities were identified within P. serraticornis, among which the cryptic species Pyrochroa bifoveata sp. n. from central Europe is described and illustrated. A comprehensive identification key to the European Pyrochroinae is also provided. Our results also suggested an historical survival of P. coccinea and P. s. kiesenwetteri in glacial refugia in Italy, and a subsequent post-glacial spread of the former species throughout the Peninsula. In contrast, the current distribution of P. s. serraticornis likely originated from a post-glacial colonization of western European relict populations, while the survival of P. bifoveata plausibly occurred in more eastern glacial refugia (e.g. Carpathian or Balkan regions). Similarly, the European populations of P. coccinea could have originated from relict populations in glacial refugia out from the Italian Peninsula. More comprehensive data on the taxonomy, ecology and biogeography of Pyrochroa are needed to learn more about these species and to help preserve the European saproxylic fauna

Pyrochroa coccinea

Pyrochroa coccinea (Linnaeus) (Figs 1A, 2A, C, E, 3 A–D, 4A, C, 5A) Diagnosis. The most diagnostic characters are structures associated with the highly sclerotized urogomphal plate: lateral lobes rounded; long, slender, straight, subparallel urogomphi; urogomphal lip ventrally furrowed and two urogomphal pits with parallel rugulae. Other diagnostic characters vs. P. serraticornis are shown in Table 1. Description of mature larva. Body length 3.5 cm (from mesal labral apex to apices of urogomphi) and maximum width 5.1 mm (across widest portion of urite VIII) (Fig. 1A). Body orthosomatic with sides subparallel, moderately sclerotized except most of cephalic capsule, mandibles, and urogomphal plate which are more heavily sclerotized; body vestiture consisting of short to moderately elongate, scattered setae. Thoracic and abdominal terga lacking parabasal ridges. Head and body dark yellowish to amber, melanisation much darker in areas of heavy sclerotization such as mandibles, urogomphi, urogomphal lip and urogomphal pits. Head. Prognathous, flattened, exserted from prothorax, about 4.5 mm width (Figs 2A, C). Epicranial suture lyriform with stem short, frontal arms complete nearly to antennal insertions, endocarinae absent (Fig. 2A). Free, symmetrical trilobate labrum, central lobe with a median thin notch, lateral lobes with high concentration of stout setae (Fig. 2A). Two pairs of stemmata on each lateral side. Antennal insertions fully exposed; antennae moderately long, slender, 3-segmented; antennomere I slightly curved outwards, narrower in the middle area, wider apically; sensorium of segment 2 small, conical; antennomere III narrower than I and II, as long as II, acutely rounded apically; setae distributed only on the inner surface of I, well distributed on all the surface of II and III. Mouthparts retracted (Figs 2 A, C). Mandibles heavily sclerotized, movable, asymmetrical, molar area of mandibles well developed; left mandible bigger, bearing a blunt molar tooth, apex rounded bidentate with two smaller subapical teeth, inner surface slightly concave (Figs 3B, C); apex of right mandible sub-securiform, inner surface heavily concave so as to make it appear bidentate from a ventral view (Figs 3A, D). Maxillae (Fig. 2E) each with 1-segmented cardo which is diagonally folded upward upon itself toward the stipes and thus appearing 2-segmented; a well-developed, undivided, pad-like maxillary articulating area; ventral surface of stipes quite glabrous, bearing few scattered setae; galea and lacinia fused to form maxillary mala; mala bearing stout apical and adoral spiniform setae and a welldeveloped pointed uncus at apico-adoral margin; 3-segmented, filiform maxillary palpus, palpomere II about 1.4X length of I, III subequal in length to II; I–II bearing stout long spiniform setae, III tapering distally, acutely rounded apically. Labium with mentum trapezoidal (Fig. 2C); submentum with a median pair of stout setae, shape elongate with sub-basal hourglass-shaped narrowing, apical margin slightly more heavily sclerotized (Fig. 2C); ligula well developed, elongate, with several pointed setae on each side in the direction of the labial palps; each labial palpus short, 2-segmented, I twice as long as II. Hypostomal rods (Fig. 2C) well developed, divergent; gular sutures separate (Fig. 2C). Thorax and Abdomen. Thorax flattened, with sides of prothorax subparallel, meso- and metathorax rounded (Fig. 1A); cervicosternum divided into three plates (Fig. 2C). Legs well developed, moderately short, 5-segmented including tarsungulus, vestiture consisting of sparse, short setae. Abdomen flattened, with sides slightly converging forward, moderately sclerotized; tergites I–VII subequal in length and width; tergite VIII approximately 2 times as long as others (Fig. 1A). Sternite VIII emarginate apically (Fig. 4C). Ventrolateral margins of abdominal laterotergite VIII emarginate, with lanceolate shape ending with more sclerotized acute apex (Fig. 4C). Tergite IX divided into four plates (Fig. 4A), hinged, capable of considerable dorso-longitudinal movement, extending ventrally, thus forming the urogomphal plate, widest basally where it forms well developed rounded lateral lobes (Figs 4A, C); surface of urogomphal plate bearing numerous, well-developed, callosities and several setigerous calli, in particular, long setae are associated with three pairs of calli on the dorsolateral surfaces (Fig. 4A), one pair of calli at the base of urogomphi and one pair of calli on the ventrolateral inner surface of urogomphi (Fig. 5A); urogomphi heavily sclerotized, long, slender, straight, subparallel, tapering and acuminate apically; ventral surface of urogomphal plate sharply excavate basally at articulation with sternites IX and X, excavation narrowing distally to bases of urogomphi and urogomphal lip, almost to form a tube under the urogomphal lip (Fig. 4C). Urogomphal plate possessing a heavily sclerotized urogomphal lip ventrally (Figs 4A, C, 5A), between the two, heavily sclerotized urogomphal pits (Fig. 5A), which, in turn, arise distally between the heavily sclerotized fixed urogomphi, and bear characteristic parallel rugulae. Sternite IX broadly transversely U-shaped (Fig. 4C), slightly invaginate in correspondence of a middle, thin, weak unsclerotized line, partially recessed into shallow emargination of sternite VIII, possessing continuous semi-circular arch of approximately 34–36 well-developed asperities along anterior margin; ventrolateral margin heavily sclerotized forming a blunt tooth (Fig. 4C). Segment X reduced, transversely ovate, with basal margin rounded, recessed into emarginations of sternite IX, visible ventrally (Fig. 4C). Spiracles. One pair of well-developed ovate thoracic spiracles ventrolaterally positioned on laterotergites along anterior margin of mesothorax. Paired, ovate abdominal spiracles, subequal in size, located on dorsolateral margin of abdominal tergite I (Fig. 1A) and ventrolateral margins of abdominal laterotergites II–VII; paired spiracles of abdominal laterotergite VIII annular-ovate, located ventrolaterally at distal 1/3 of its length (Fig. 4C).Published as part of Molfini, Marco, Giulio, Andrea Di, Mancini, Emiliano & Bologna, Marco A., 2021, Larval features illuminating adult taxonomy? Case study in the European cardinal beetle species of the genus Pyrochroa (Coleoptera: Pyrochroidae: Pyrochroinae), pp. 337-348 in Zootaxa 4966 (3) on pages 339-341, DOI: 10.11646/zootaxa.4966.3.5, http://zenodo.org/record/473673

Pyrochroa serraticornis

Pyrochroa serraticornis (Scopoli) (Figs 1B, 2B, D, F, 3E–H, 4B, D, 5B) Diagnosis. The most diagnostic characters are structures associated with the highly sclerotized urogomphal plate: lateral lobes acute rounded; sclerotized conical teeth between lateral lobes and the straight parallel urogomphi; urogomphal lip flat, ventrally to the single urogomphal pit with parallel rugulae. Other diagnostic characters vs. P. coccinea are shown in Table 1. Description of mature larva. Body length about 2.4 cm (from mesal labral apex to apices of urogomphi) and maximum width about 3.8 mm (across widest portion of urite VIII) (Fig. 1B). Body orthosomatic with sides subparallel, moderately sclerotized except much of cephalic capsule, mandibles, and urogomphal plate more heavily sclerotized; body vestiture consisting of short to moderately elongate, scattered setae. Thoracic and abdominal tergites II–VII and IX lacking parabasal ridges; abdominal tergites I and VIII with weakly formed parabasal ridge (Figs 1B, 4B). Head and body from light yellowish to darker, melanisation much darker in areas of heavy sclerotization such as mandibles, urogomphi, urogomphal lip and urogomphal pit. Head. Prognathous, flattened, exserted from prothorax, about 3.3 mm width (Figs 2B, D). Epicranial suture lyriform with stem short, frontal arms complete nearly to antennal insertions, endocarinae absent (Fig. 2B). Free, symmetrical trilobate labrum with scattered slender setae (Fig. 2B). Two pair of stemmata on each lateral side, parallel with antennal insertions. Antennal insertions fully exposed; antennae moderately long, stout, 3-segmented; antennomere I quite straight, narrower in the middle area, wider apically; sensorium of II small, drop-shaped; antennomere III narrower than I–II, about 2/3 the length of II, acutely rounded apically; setae well distributed on all the surface of antennomers. Mouthparts retracted (Figs 2B, D). Mandibles heavily sclerotized, movable, asymmetrical, molar area of mandibles well developed; left mandible bigger, bearing a prominent, hooked inwards, molar tooth, apex sub-securiforme with one small subapical expansion; inner surface bearing two longitudinal depressions so as to make it appear tridentate from a ventral view (Fig. 3F, G); apex of right mandible securiforme, inner surface concave so as to make it appear bidentate from a ventral view (Figs 3E, H). Maxillae (Fig. 2F) each with 1-segmented cardo which is diagonally folded upward upon itself toward the stipes and thus appearing 2- segmented; a well-developed, undivided, pad-like maxillary articulating area; ventral surface of stipes bearing few scattered spiniform setae; galea and lacinia fused to form maxillary mala; mala bearing stout apical and adoral spiniform setae and a well-developed pointed uncus at apico-adoral margin; 3-segmented, filiform maxillary palpus, palpomere II about 1.2 times as long as I, palpomere III subequal in length to II, tapering distally, acutely rounded apically. Labium with mentum trapezoidal (Fig. 2D); submentum glabrous, shape elongate with sides shallowing sinuate basally, apical margin slightly more heavily sclerotized (Fig. 2D); ligula well developed, elongate, almost glabrous, bearing few setae hardly characterized from our samples; each labial palpus short, 2-segmented, palpomere I twice as long as II. Hypostomal rods (Fig. 2D) well developed, divergent; gular sutures separate (Fig. 2D). Thorax and Abdomen. Thorax flattened, segmentation well developed, sides of prothorax subparallel, meso- and metathorax rounded (Fig. 1B); cervicosternum divided into three plates (Fig. 2D). Legs well developed, moderately short, 5-segmented including tarsungulus, vestiture consisting of sparse, short setae.Abdomen flattened, sides slightly converging forward, moderately sclerotized, tergites I–VII subequal in length and width; tergite VIII approximately 2.8 as long as each other (Fig. 1B). Sternite VIII emarginate apically (Fig. 4D). Ventrolateral margins of abdominal laterotergite VIII emarginate, with lanceolate shape, ending with more sclerotized acute apex forming a lateral tooth visible even from dorsal view (Figs 1B, 4D). Tergite IX divided into four plates (Fig. 4B), hinged, capable of considerable dorso-longitudinal movement, extending ventrally, thus forming the urogomphal plate, widest basally where it forms well developed acute rounded lateral lobes (Figs 4B, D); surface of urogomphal plate bearing numerous, well-developed, callosities and several setigerous calli, in particular, long setae are associated with three pair of calli on the dorsolateral surfaces and one pair of calli at the base of urogomphi (Fig. 4B); a pair of well developed, highly sclerotized conical teeth between urogomphi and lateral lobes (Figs 4B, D); urogomphi heavily sclerotized, long, slender, straight, parallel, tapering and acuminate apically; ventral surface of urogomphal plate sharply excavate basally at articulation with sternites IX–X, excavation narrowing distally to bases of urogomphi and urogomphal lip forming a thin slot (Fig. 4D). Urogomphal plate possessing a heavily sclerotized urogomphal lip ventrally (Figs 4B, D, 5B), forming the ventral surface of the wide heavily sclerotized single urogomphal pit (Fig. 5B), which fills the space between the heavily sclerotized fixed urogomphi, and bear characteristic parallel rugulae. Sternite IX broadly transversely U-shaped (Fig. 4D), partially recessed into shallow emargination of sternite VIII, possessing continuous semicircular arch of approximately 32 well-developed asperities along anterior margin; ventrolateral margin heavily sclerotized forming an acute tooth slightly visible even in dorsal view (Figs 1B, 4D). Segment X reduced, transversely ovate, basal margin rounded, recessed into emarginations of sternite IX, visible ventrally (Fig. 4D). Spiracles. One pair of well-developed, ovate, thoracic spiracles situated ventrolaterally on laterotergite along anterior end of mesothorax. Paired, sub-ovate abdominal spiracles, subequal in size, located on dorsolateral margin of abdominal tergite I (Fig. 1B) and ventrolateral margins of abdominal laterotergites II–VII; paired spiracles of abdominal laterotergite VIII annular-ovate, located ventrolaterally at distal 1/3 of its length (Fig. 4D).Published as part of Molfini, Marco, Giulio, Andrea Di, Mancini, Emiliano & Bologna, Marco A., 2021, Larval features illuminating adult taxonomy? Case study in the European cardinal beetle species of the genus Pyrochroa (Coleoptera: Pyrochroidae: Pyrochroinae), pp. 337-348 in Zootaxa 4966 (3) on pages 341-345, DOI: 10.11646/zootaxa.4966.3.5, http://zenodo.org/record/473673

Is Lycorma delicatula (Hemiptera: Fulgoridae) a blooming threat to citrus?

Examining the host range of emerging invasive insects is essential to assess their invasion potential and to anticipate the negative impacts of their spread. The ongoing North American invasion of spotted lanternfly (SLF) [Lycorma delicatula (White, 1845)] threatens agricultural, urban, and natural areas. The survival and development of SLF nymphs on Washington navel orange [Citrus sinensis (L.) Osbeck (Sapindales: Rutaceae)] trees were assessed in a quarantine facility. Results indicated that SLF nymphs can develop to at least the third instar by feeding exclusively on Washington navel orange. This finding suggests that, at least up to the third stage of nymphal development, Washington navel orange might be a suitable host for SLF, highlighting the possibility that this invasive pest represents an unrecognized threat to this globally important crop and possibly to other Citrus species.This work was supported, in part, by California Department of Food and Agriculture’s (CDFA) Plant Health and Pest Prevention Services Division (PHPPS) under award number A21-2688-S004 to UC Riverside. Additional support was provided by CDFA’s Office of Environmental Farming and Innovation, Proactive Integrated Pest Management Solutions Grant Program, award number 18-0632-000-SG “Proactive biological control of spotted lanternfly, Lycorma delicatula (Hemiptera: Fulgoridae)”.info:eu-repo/semantics/publishedVersio

Strange loves: a remarkable case of aberrant copulation in beetles (Coleoptera: Meloidae, Chrysomelidae)

A case of copulation between two mimic and repellent beetle species (a male of Timarcha fracassii, and a female of Meloe autumnalis), belonging to distinct families (Chrysomelidae, Meloidae), is recorded

Microdon Meigen 1803

Keys to immature stages of the European species of Microdon First instar larvae 1 Body about 1.5 times as long as wide; posterior spiracular tubercle long (more than 1.5 times as long as wide at base); marginal band elongate, showing eight waves on each side...................................... M. myrmicae / M. mutabilis - Body about two times as long as wide; posterior spiracular tubercle very short (wider than long at base); marginal band short, showing 10 waves on each side................................................................... M. analis Puparia 1 Most of the dorsal surface of puparium smooth and bare, with reticulation reduced to a narrow, lateral belt.............. 2 - Dorsal surface of puparium completely covered by a dorsal reticulation.......................................... 3 2 Anterior spiracular tubercles conical, tapering at apex; anterior spiracular tubercles about 1.4 times as long as wide at base; each reticulation process composed of groups of umbrella-like structures................................... M. myrmicae - Anterior spiracular tubercles dome-shaped, blunt at apex; anterior spiracular tubercles about 0.8 times as long as wide at base; each reticulation process composed of stringy, extended projections.................................... M. mutabilis 3 Meshes of the dorsal reticulation no broader than the basal diameter of the posterior spiracular tubercle; posterior spiracular tubercle wider than long; anterior spiracular tubercles straight................................................. 4 - Meshes of the dorsal reticulation two times as broad as the basal diameter of the posterior spiracular tubercle; posterior spiracular tubercle longer than wide; anterior spilacural tubercles laterally curved................................. M. devius 4 Anterior spiracular tubercles equal or more than two times as long as wide....................................... 5 - Anterior spiracular tubercles one time only as long as wide, or shorter than wide.............................. M. miki 5 Anterior spiracular tubercles nearly cylindrical, about three times as long as wide; posterior spiracular tubercle light-brown contrasting with the reddish-brown apical spiracular plates.............................................. M. analis - Anterior spiracular tubercles clearly conical, about two times as long as wide; posterior spiracular tubercle uniformly reddishbrown....................................................................................... M. majorPublished as part of Scarparo, Giulia, Wolton, Robert, Molfini, Marco, Pinna, Luigi Cao & Ulio, Andrea Di Gi-, 2020, Comparative morphology of myrmecophilous immature stages of European Microdon species (Diptera: Syrphidae): updated identification key and new diagnostic characters, pp. 348-370 in Zootaxa 4789 (2) on pages 364-365, DOI: 10.11646/zootaxa.4789.2.2, http://zenodo.org/record/399083

Microdon myrmicae Schonrogge

Immature stages of Microdon myrmicae Egg Figs 1, 11 Width = 587.4 ± 36.61 µm; length = 1.08 ± 0.03 mm (n = 10). First instar larva Figs 2−6, 11, Supplementary Material 1 Body width = 679.5 ± 55.42 µm; body length = 0.996 ± 0.08 mm (n = 10). Body features. Body with regularly rounded sides. Four longitudinal grooves present dorsally (Fig. 2A) dividing dorsal body surface into five main longitudinal fields: one medial, two lateral and two external fields. Medial field partially divided into two halves by a longitudinal, medial line. Dorsal surface with regularly spaced “flower-like” sensilla (Figs 3A, E): medial field with two longitudinal rows of nine sensilla; each lateral field with 13 sensilla arranged in two rows (seven along lateral groove and six along medial groove); each external field with one row of 10 sensilla. Each sensillum (Fig. 3E) composed of a cylindrical base, with many imbricate, thick sculpticels, apically with a medial flower-like structure with a variable number (5–10) of long lobes, pointed at tip, encircling a medial dome. Ventral surface covered medially by pointed microsculpture, finely pilose on sides. Ventral flower-like sensilla (Figs 3C, G) similar to dorsal ones except for flat, soft, unsculptured base and flat, thin, distinctly pointed lobes. Pseudocephalon. Two pairs of sensorial organs on dorsal surface of pseudocephalon (Figs 4E, G), one anterior (Fig. 4E) and one posterior (Fig. 4G), each composed of clusters of four short and one long trichoid sensilla emerging from bulbous, hollow base. Posterior spiracular tubercle. Impair respiratory structure, elongated in shape; surface of spiracular tubercle with peculiar microsculpture, completely covered by imbricate, sclerotised scales with an irregularly indented apex (Fig. 5A). Apex of tubercle with two circular smooth plates, slightly convex (Fig. 5C). Marginal band. Distinctly long fringe, length of processes regularly varying, showing eight waves on each side (Figs 2A, E, 6A, C, E). Each individual process composed of an elongate stem and an apical fringed brush (Figs 6A, C, E); the stem showing two very different surfaces: dorsal surface apparently articulated with 4–5 imbricated joints, the distal one fringed apically (Figs 6A, E); ventral surface completely smooth (Fig. 6C). Third instar larva Figs 7−9, 11 Body width = 4.5 ± 0.7 mm; body length = 6.1 ± 0.8 mm (n = 10). Dorsal reticulation. Dorsal reticulation reduced to a narrow, lateral belt around the perimeter of abdomen (Figs 7A, B). Each reticulation process showing sub circular groups of 5–9 umbrella-like structures with a flattened, circular apex (Figs 7A, B). Posterior spiracular tubercle. Dome shaped with two round holes spaced 1.2 times as long as their diameter; apex divided into two halves (Figs 8A, B) by a narrow furrow showing irregular plates. Marginal band. Processes on the marginal band short and thick, set close to one another, parallel, radially projecting, with suboval basal “joints”, not imbricated, the last one produced into a flat medial brush. Processes on the marginal band of three types (Figs 9A, B): type one basally 4–jointed and apically single and flat, with medial brush 2–lobed; type two basally 3–jointed and apically bifurcate, with medial brush 1–lobed; type three basally 3–jointed and apically produced into a group of three spiniform setae. Type one and type two regularly alternating in sequence, type three irregularly present between two type one processes. Puparium Figs 10, 11 Body width = 6.1 ± 0.5 mm; body length = 7.9 ± 0.7 mm (n = 10). Anterior spiracular tubercles. Length of each tubercle about 1.4 times as long as wide, conical, tapering at the apex (Fig. 10A, B), smooth at the base, with the apex furrowed by about 150 respiratory fissures (Figs 10A); each fissure laying on a small papilla (Fig. 10B).Published as part of Scarparo, Giulia, Wolton, Robert, Molfini, Marco, Pinna, Luigi Cao & Ulio, Andrea Di Gi-, 2020, Comparative morphology of myrmecophilous immature stages of European Microdon species (Diptera: Syrphidae): updated identification key and new diagnostic characters, pp. 348-370 in Zootaxa 4789 (2) on pages 354-357, DOI: 10.11646/zootaxa.4789.2.2, http://zenodo.org/record/399083