Monograph
Print
Monograph
The world woodlouse flies (Diptera, Rhinophoridae)
expand article infoPierfilippo Cerretti, Davide Badano§, Silvia Gisondi|, Giuseppe Lo Giudice#, Thomas Pape|
‡ Sapienza University of Rome, Rome, Italy
§ DISTAV, University of Genoa, Genoa, Italy
| University of Copenhagen, Copenhagen, Denmark
¶ CUFA, Raggruppamento Carabinieri Biodiversità, Verona, Italy
# LaNaBIT – Laboratorio Nazionale Tassonomia e Biomonitoraggio Invertebrati, Verona, Italy
Open Access

Abstract

The world Rhinophoridae are catalogued, recognising 33 genera and 177 species. Nomenclatural information is provided for all genus-group and species-group names, including lists of synonyms and name-bearing type data. Species distributions are recorded by country. A key to the world genera is presented. Four new genera are erected to accommodate five new species, which do not fit within any of the current generic concepts in Rhinophoridae, according to the results of a morphology-based phylogenetic analysis: Marshallicona Cerretti & Pape with type species Marshallicona quitu Cerretti & Pape, gen. et sp. nov. (Ecuador); Maurhinophora Cerretti & Pape with type species Maurhinophora indoceanica Cerretti & Pape, gen. et sp. nov. (Mauritius); Neotarsina Cerretti & Pape with type species Neotarsina caraibica Cerretti & Pape, gen. et sp. nov. (Trinidad and Tobago) and Neotarsina andina Cerretti & Pape, sp. nov. (Peru); Kinabalumyia Cerretti & Pape with type species Kinabalumyia pinax Cerretti & Pape, gen. et sp. nov. (Malaysia, Sabah). The genus Aporeomyia Pape & Shima (type species Aporeomyia antennalis Pape & Shima), originally assigned to Tachinidae, is here reassigned to Rhinophoridae based on a reassessment of the homologies of the male terminalia. The following five species-group names, which were previously treated as junior synonyms or nomina dubia, are recognised as valid species names: Acompomintho caucasica (Villeneuve, 1908), stat. rev. [from nomen dubium to valid species]; Acompomintho sinensis (Villeneuve, 1936), stat. rev. [from nomen dubium to valid species]; Stevenia bertei (Rondani, 1865), stat. rev. [from nomen dubium to valid species]; Stevenia sardoa Villeneuve, 1920, stat. rev. [from junior synonym of Rhinophora deceptoria Loew, 1847 to valid species]; Stevenia subalbida (Villeneuve, 1911), stat. rev. [from junior synonym of Rhinophora deceptoria Loew, 1847 to valid species]. Reversal of precedence is invoked for the following case of subjective synonymy to promote stability in nomenclature: Rhinophora lepida (Meigen, 1824), nomen protectum, and Musca parcus Harris, 1780: 144, nomen oblitum. New generic and specific synonymies are proposed for the following two names: Mimodexia Rohdendorf, 1935, junior synonym of Tromodesia Rondani, 1856, syn. nov. and Ptilocheta tacchetti Rondani, 1865, junior synonym of Stevenia obscuripennis (Loew, 1847), syn. nov. The following new combinations are proposed: Acompomintho sinensis (Villeneuve, 1936), comb. nov. [transferred from Tricogena Robineau-Desvoidy, 1830]; Tromodesia guzari (Rohdendorf, 1935), comb. nov. [transferred from Mimodexia Rohdendorf, 1935]; Tromodesia intermedia (Rohdendorf, 1935), comb. nov. [transferred from Mimodexia Rohdendorf, 1935]; Tromodesia lindneriana (Rohdendorf, 1961), comb. nov. [transferred from Mimodexia Rohdendorf, 1935]; Tromodesia magnifica (Rohdendorf, 1935), comb. nov. [transferred from Mimodexia Rohdendorf, 1935]; Tromodesia obscurior (Rohdendorf, 1935), comb. nov. [transferred from Mimodexia Rohdendorf, 1935]; Tromodesia pallidissima (Rohdendorf, 1935), comb. nov. [transferred from Mimodexia Rohdendorf, 1935]; Tromodesia setiventris (Rohdendorf, 1935), comb. nov. [transferred from Mimodexia Rohdendorf, 1935] and Tromodesia shachrudi (Rohdendorf, 1935), comb. nov. [transferred from Mimodexia Rohdendorf, 1935].

Keywords

Catalogue, cladistic analysis, classification, distribution, new taxa, Oestroidea, parasitoids, systematics, zoological nomenclature

Introduction

Rhinophoridae are a small oestroid family with 33 genera and 177 species, recognised as of the present catalogue. The family was earlier considered by several authors to be entirely of Old World distribution and including very few native species outside the Palaearctic Region. This notion was at least partly due to the then superficially sampled tropical and southern hemisphere subtropical faunas, which are continuously revealing new taxa (Crosskey 1977, Colless 1994, Pape 1998a, Pape and Arnaud 2001, Cerretti and Pape 2012, Cerretti et al. 2014a).

Adult Rhinophoridae present no unique autapomorphies (Fig. 1), thus they are difficult to key out in conventional family-level keys (e.g., Crosskey 1977, Marshall 2012). However, their larvae are highly specialised woodlouse parasitoids and, to the extent they are known, provide unambiguous evidence for family affiliation and monophyly (Bedding 1973, Pape and Arnaud 2001, Cerretti et al. 2014a). The woodlouse-parasitising habit is unique not only within Diptera but within all Insecta. The distribution of Rhinophoridae largely matches that of woodlice, and the peak of rhinophorid diversity and abundance appears to be the Turano-Mediterranean area of the western Palaearctic (see below).

Figure 1. 

Rhinophoridae photographed in nature. A Bezzimyia sp. (Ecuador) B Bixinia winkleri (Australia) C Marshallicona quitu (Ecuador) D Paykullia maculata (Finland) E Stevenia sp. (Italy) F Tricogena rubricosa (Finland) G Phyto sp. (Italy). Photographs by Steve Marshall (A, C), P.C. (B, E, G), Håkon Haraldseide (D, F).

The family Rhinophoridae is a member of the Oestroidea, but its phylogenetic position within this clade is still unresolved (Pape 1992, Rognes 1997, Pape and Arnaud 2001, Kutty et al. 2010, Winkler et al. 2015, Cerretti et al. 2017, Cerretti et al. 2019, Stireman et al. 2019, Kutty et al. 2019). The most recent analyses based on morphology indicate a sister-group relationship to Rhiniidae (Pape and Arnaud 2001) or to the monotypic New Zealand family Mystacinobiidae, with Rhiniidae subordinate to these (Cerretti et al. 2017), which find most arguments from first instar larval morphology, which is very superficially known for the latter group (Ferrar 1987, Rognes 2002). Interestingly, the most recent analyses based on molecular data retrieve Rhiniidae as sister to Bengaliinae (Cerretti et al. 2017, Stireman et al. 2019, Kutty et al. 2019, Cerretti et al. 2019), and the extensive transcriptome data of Kutty et al. (2019) point to a phylogenetic grouping of Rhinophoridae with the macrolarviparous Helicoboscinae and Ameniinae, which is unexpected from a morphological as well as a biological point of view. The phylogenetic position of Rhinophoridae within Oestroidea remains ambiguous.

Peris and González-Mora (2007) compiled the first world catalogue for the family Rhinophoridae, but with insufficient distributional data and without data on type localities and types. Taking into account the substantial number of genera and species recently described from outside the Palaearctic, which have not been included in any regional catalogue, the significant recent exclusions (i.e., Alvamaja chlorometallica Rognes and the five species of the Phyto carinata species group now assigned to the genus Morinia Robineau-Desvoidy, see Cerretti et al. 2019) as well as a new inclusion (Aporeomyia Shima and Pape, as proposed herein) of taxa, we consider it both helpful and timely to provide a fully updated world catalogue of the entire family. We also take this opportunity to describe four new genera to accommodate five new species, produce a key to all genera, and perform a genus-level phylogenetic analysis of the Rhinophoridae in order to support the newly proposed genera in a cladistic framework.

Materials and methods

Systematics

Taxa, specimens

All genera were studied, based on an extensive representation of the included species, in order to construct a key to the genera of the world and perform a genus-level phylogenetic analysis. Dissections of male terminalia were performed according to the procedure described by Cerretti and Shima (2011). Briefly, this procedure involves the removal of the abdomen, partial clearing in 10% KOH, dissection and rinsing of terminalia, reattachment of the abdomen to the specimen, and final storage of the terminalia in a glycerine-filled microvial pinned with the source specimen. Figures 2 and 3 summarise the morphological terminology of the adults (except terminalia) and the measurements applied in the present work. Morphological terminology of terminalia (both male and female) and preimaginal instars follow Cumming and Wood (2017) and Pape and Arnaud (2001), respectively.

Figure 2. 

Morphological terminology of Rhinophoridae, head, thorax and abdomen. A Chaetotaxy of head, thorax and abdomen in dorsal view B, C arrangement of setae of postpronotum: B three setae arranged in shallow triangle C three setae standing in a nearly right-angled triangle D thorax, sclerites and chaetotaxy, in lateral view E wing veins and cells. Drawings by Giulia Bellanti.

Figure 3. 

Morphological terminology of Rhinophoridae, head. A Standard measurements in dorsal view B head structures in frontal view C head structures in lateral view D standard measurements in lateral view. Drawings by Giulia Bellanti.

Data from each type specimen are given verbatim, with information for each line separated by a slash (/) ; in cases with more than one label, these are separated with a double slash (//). Additional information of relevance, but not appearing on the label(s), is given in brackets. Finally, the acronym of the repository is cited in parentheses.

Photography, SEM

Photographic images of habitus, head, wing, legs, abdomen and male terminalia were produced using a MZ 12.5 stereoscopic microscope (Leica, Germany) equipped with a DS-L1 Nikon digital camera (Nikon, Tokyo). Photographic images of terminalia were produced using a DM LS microscope (Leica, Germany) equipped with the camera described above. Focus stacking with the image stacking software CombineZM (Hadley, UK) was used to merge 15–45 photos of each specimen/structure, taken at different focal planes, into high-resolution images. Additional images were produced with a Hitachi TM1000 environmental scanning electron microscope (ESEM) from uncoated pinned specimens, using Adobe Photoshop for colouration of specific structural details.

Phylogenetic analysis

We adapted the data matrix of Cerretti et al. (2014a) to assess the most probable phylogenetic affinities and associated support of the newly proposed genera. The matrix consists of 99 morphological characters: 82 from adult stage (1–52: body of both sexes, except terminalia; 53–81: male terminalia; 82: female terminalia), 17 from first instar larva (Table 1); rhinophorid diversity is represented by a sample of 57 species (comprising at least one species for each genus), plus Musca spp. (Muscidae) and a selection of oestroid taxa as outgroups (Table 2; the nexus format of the data matrix is available from MorphoBank Project 3576). In the case of Kinabalumyia pinax gen. et sp. nov., described in the present paper based on two males from Sabah (Malaysia), female characters were scored from a female from Palawan (Philippines), which is here considered congeneric but is left unidentified (Kinabalumyia sp. 1, see below); the two sets of characters were concatenated and treated as a single operational taxonomic unit (OTU) in all our analyses, which is shown in the tree under the name of Kinabalumyia pinax for practical reasons.

Table 1.

List of characters and states.

ADULT (body parts, except terminalia)
1 Arista, microtrichia length: (0) shorter than maximum diameter of arista; (1) distinctly longer.
2 Male arista, microtrichia: (0) not bottlebrush-like; (1) bottlebrush-like.
3 Arista, development: (0) normally developed; (1) shortened.
4 Arista, setae: (0) absent; (1) present.
5 Arista, thickening: (0) at most on basal fourth; (1) at least on basal 3/4.
6 First aristomere, length: (0) short, at most as long as wide; (1) approx. 2–3 times as long as wide; (2) at least 4 times as long as wide.
7 Lunule, setae: (0) bare; (1) with setae.
8 Female, groove between fronto-orbital plate and parafacial: (0) absent; (1) present.
9 Male, proclinate orbital setae: (0) absent; (1) present, at least as long as frontal setae; (2) present, distinctly shorter than frontal setae.
10 Female, proclinate orbital setae: (0) absent; (1) present, at least as long as frontal setae; (2) present, distinctly shorter than frontal setae.
11 Proclinate orbital setae, when present, number: (0) one or two; (1) more than two.
12 Vibrissal triangle: (0) normal; (1) projected.
13 Shape of lower facial margin, shape: (0) not sunken; (1) deeply sunken (i.e., concave).
14 Facial plate, shape: (0) not sunken; (1) deeply sunken.
15 Facial plate, median carina: (0) absent; (1) present.
16 Compound eye, posterior margin: (0) not indented; (1) indented.
17 Parafacial, setosity: (0) bare in ventral half; (1) setose in ventral half.
18 Parafacial setae, configuration: (0) short, scattered and proclinate; (1) long, robust and medioclinate.
19 Frontal setae, orientation of dorsal-most pair: (0) converging in the middle and crossed; (1) sub-parallel.
20 Mouthparts, development: (0) normally developed (i.e., prementum between 2 and 6 times as long as wide); (1) both prementum and labella strongly reduced (vestigial).
21 Palpus: (0) normally developed; (1) reduced; (2) absent.
22 Occiput, setae: (0) black, normal; (1) pale and hair-like.
23 Postpronotal setae, number: (0) four (or more) setae; (1) three setae; (2) two setae; (3) one seta; (4) none.
24 Postpronotal setae, if three or more, position: (0) mid-basal seta in line, or nearly so, with inner and outer basal setae; (1) mid-basal seta displaced anteriorly to line between inner and outer basal setae (i.e., three robust, basal setae in an almost right-angled triangle).
25 First postsutural supra-alar seta, size: (0) present, approx. as long as notopleural setae; (1) absent or very short and hair-like; (2) strong, distinctly longer and thicker than notopleural setae.
26 Subscutellum, development: (0) flat or concave; (1) moderately swollen; (2) strongly swollen.
27 Subscutellum, sclerotisation: (0) not fully sclerotised; (1) entirely sclerotised.
28 Scutellum, apical setae: (0) present; (1) absent.
29 Katepimeron, setae: (0) with at least one seta anteriorly; (1) entirely bare.
30 Metathoracic spiracular lappets: (0) practically absent; (1) small, sub-equal in size and directed outwards; (2) unequal in size (posterior one distinctly larger) and both lappets closing the spiracle like an operculum.
31 Male fore tarsus: (0) normal; (1) laterally compressed.
32 Female fore tarsus: (0) normal; (1) laterally compressed.
33 Shape of tibiae of mid and hind legs: (0) normal; (1) laterally compressed and distinctly keeled dorsally.
34 Leg chaetotaxy: (0) not particularly modified; (1) reduced and with almost no (strong) setae.
35 Hind coxa, posterodorsal margin: (0) bare; (1) with (1–4) setae.
36 Lower calypter, shape: (0) semi-circular; (1) tongue-shaped.
37 Long trichia along margin of lower calypter: (0) absent; (1) present.
38 Female wing, development: (0) normally developed; (1) brachypterous; (2) micropterous.
39 Costa, indentation at level of R4+5: (0) absent; (1) present.
40 Male wing pattern, three distinct whitish spots: (0) absent; (1) present.
41 Female wing pattern, posterodistal whitish spot: (0) absent; (1) present.
42 Vein R1, dorsal setae: (0) absent; (1) with a row of setulae along whole length; (2) with 1–10 setae at most on apical fourth.
43 Vein R4+5, dorsal setae: (0) present; (1) absent.
44 Vein R4+5, extent of dorsal setae, when present: (0) only at base; (1) reaching at least crossvein r-m;
45 Bend of vein M 1: (0) distinct; (1) not distinct.
46 Vein M 1, position of bend: (0) well removed from wing margin; (1) very close to wing margin but distinct.
47 Vein M 1, extent of distal part: (0) not vanishing on wing margin; (1) gradually vanishing on wing membrane.
48 Vein M 1, apical termination: (0) joining R4+5 so that cell r4+5 is closed; (1) joining costa, so that cell r4+5 is open.
49 Crossvein dm-m, inclination: (0) forming a right angle with proximal section of M4; (1) forming an acute angle with proximal section of M4.
50 CuA+CuP: (0) not reaching wing margin; (1) reaching wing margin.
51 Tergite 3, median marginal setae: (0) present; (1) absent or very short and recumbent.
52 Tergite 4, marginal setae: (0) a regular row of more or less erect marginal setae; (1) marginal setae absent or not differentiated from general abdominal setulae.
ADULT (male terminalia)
53 Sternite 5, posteromedian notch: (0) present; (1) absent.
54 Sternite 5, shape of transversal section: (0) U-shaped (i.e., folds up laterally); (1) almost flat.
55 Sternite 5, median tooth-like apophysis on lateral lobe: (0) absent; (1) present.
56 Tergite 6, median marginal setae: (0) present; (1) absent.
57 Tergite 6, shape: (0) normal (plate-like); (1) posteriorly indented; (2) divided into two hemitergites; (3) reduced, absent.
58 Connection between tergite 6 and syntergosternite 7+8: (0) membranous; (1) tergite 6 and syntergosternite 7+8 fused.
59 Connection between sternite 6 and syntergosternite 7+8 on right side: (0) membranous; (1) fused.
60 Cerci, shape: (0) normally developed; (1) short and sub-globular.
61 Cerci, medial connection: (0) not fused at all medially; (1) at least partly fused medially.
62 Surstylus, shape: (0) not divided; (1) with a large base distally divided into a lateral, wide, rounded lobe and into a postero-median finger-like apophysis.
63 Surstylus, bifid inner median extension: (0) absent; (1) present.
64 Surstylus, setae on median extension: (0) absent; (1) present.
65 Bacilliform sclerite and laterobasal margin of surstylus, connection: (0) articulated, not fused; (1) firmly fused.
66 Surstylus and epandrium, connection: (0) membranous; (1) surstylus and epandrium fused.
67 Hypandrial arms, shape: (0) firmly fused postero-medially, entirely encircling base of phallus; (1) more or less converging medially but not fused.
68 Phallic guide and pregonites, connection: (0) membranous (i.e., not fused); (1) sclerotised (i.e., fused).
69 Postgonite, anterior seta: (0) present; (1) absent.
70 Extension(s) of dorsal sclerite of distiphallus: (0) present; (1) absent.
71 Extension(s) of dorsal sclerite of distiphallus, longitudinal division: (0) divided into two hemisclerites or at least partly unfused medially (i.e., distally bifid); (1) entirely fused mid-dorsally into a single sclerotisation (i.e., distally not bifid).
72 Extension(s) of dorsal sclerite of distiphallus, sclerotised connection with dorsal sclerite of distiphallus: (0) present; (1) absent.
73 Membranous flag distal to extension of dorsal sclerite of distiphallus: (0) absent; (1) present.
74 Median process of ventral sclerotisation of distiphallus: (0) present; (1) absent.
75 Median process of ventral sclerotisation of distiphallus, connection with ventral plate: (0) not interrupted, running from the ventral plate to tip of phallus; (1) interrupted proximally and not connected to ventral plate.
76 Median process of ventral sclerotisation of distiphallus, longitudinal division: (0) not divided longitudinally; (1) divided longitudinally.
77 Median process of ventral sclerotisation of distiphallus, length: (0) normal; (1) very long (i.e., ending far beyond tip of acrophallus).
78 Median process of ventral sclerite of distiphallus, asymmetry: (0) absent; (1) present.
79 Acrophallus, shape: (0) simple, unmodified (with one opening); (1) distinctly tripartite (with three openings).
80 Semi-cylindrical dorsal sclerite of acrophallus: (0) absent; (1) present.
81 Distiphallus, helmet-shaped, partly sclerotised envelope: (0) absent; (1) present.
ADULT (female terminalia)
82 Ovipositor: (0) long and telescopic; (1) shortened.
LARVA (first instar)
83 Labrum: (0) well developed; (1) reduced.
84 Locomotory behaviour: (0) creeping (peristalsis); (1) leech-like; (2) somersaulting.
85 Antenna: (0) normally developed; (1) long and tapering.
86 Posterior part of anal division modified as a terminal sucker: (0) no; (1) yes.
87 Mandibles: (0) well developed; (1) reduced.
88 Shape of mandible: (0) normal, hook-like; (1) toothed or serrated.
89 Labrum, connection with cephaloskeleton: (0) not fused; (1) fused.
90 Body shape: (0) subfusiform; (1) slightly flattened dorsoventrally.
91 Parastomal bar of cephaloskeleton: (0) reduced or not elongated; (1) long and slender.
92 Longitudinal incision on parastomal bar of cephaloskeleton: (0) absent; (1) present.
93 Fleshy protuberances (= prolegs) on segments: (0) absent; (1) present.
94 Shape of ventral part of pseudocephalon: (0) normal (i.e., not elongated); (1) elongated.
95 Longitudinal cuticular ridges posteroventrally on anal division: (0) absent; (1) present.
96 Tongue-like projection posterodorsally on anal division: (0) absent; (1) present.
97 Pair of more or less elongated or globular vesicles posteroventrally on anal division: (0) absent; (1) present.
98 Shape of posteroventral vesicles, is present: (0) sub-globular; (1) long and tapering.
99 Mandible, number of teeth (if present): (0) two; (1) three or more.
Table 2.

List of material examined for the cladistic analysis. An asterisk (*) denotes taxa not included in the dataset of Cerretti et al. (2014a).

Family/subfamily Genus/species Country, State or Region [collection acronym] References
Muscidae/Muscinae Musca spp. Italy, Latium [MZUR]
Calliphoridae/Ameniinae Amenia sp. Australia, Queensland [MZUR] Crosskey (1965)
Calliphoridae/Bengaliinae Bengalia sp. Thailand, Chiang Mai [MZUR] Rognes (2011)
Calliphoridae/Calliphorinae Calliphora vomitoria (Linnaeus, 1758) Italy [MZUR] Rognes (1991)
Rognes (1997)
Calliphoridae/Helicoboscinae Eurychaeta muscaria (Meigen, 1826) Italy, Latium and Veneto [MZUR] Rognes (1986)
Rognes 1991)
Calliphoridae/Luciliinae Lucilia sericata (Meigen, 1826) Italy, Sardinia [MZUR] Rognes (1991)
Calliphoridae/Melanomyiinae Melinda gentilis Robineau-Desvoidy, 1830 Italy, Sardinia [MZUR] Rognes (1991)
Oestridae/Cuterebrinae Cuterebra austeni Sabrosky, 1986 USA, New Mexico [MZUR]
Polleniidae Pollenia paupera Rondani, 1862 Italy, Latium [MZUR]
Rhiniidae Rhyncomya impavida (Rossi, 1790) Italy, Latium [MZUR]
Tachinidae/Tachininae Macquartia tenebricosa (Meigen, 1824) Italy, Latium [MZUR]
Rhinophoridae Acompomintho lobata* Japan, Fukuoka [NHMD] Pape (1986); Kato and Tachi (2016)
Apomorphyto inbio Costa Rica, Guanacaste [INBio]
Aporeomyia sp.* Malaysia, Sabah [CNC, NHMD] Pape and Shima (1993)
Axinia disjuncta Australia, Queensland [ANIC] Colless (1994)
Axinia lucaris Australia, Queensland [NHRS] Colless (1994)
Axinia miranda Papua New Guinea [ANIC] Colless (1994)
Azaisia sp.* Portugal, Madeira [NHMD]
Baniassa fascipennis Israel [TAU]
Bezzimyia barbarista Costa Rica, Alajuela [NHMD] Pape and Arnaud (2001)
Bezzimyia bisecta Costa Rica: Monteverde and Guanacaste [NHMD] Pape and Arnaud (2001)
Pape 2010)
Bezzimyia busckii Costa Rica, Guanacaste [NHMD] Pape and Arnaud (2001)
Pape (2010)
Bezzimyia hansoni Costa Rica, Limón [NHMD] Pape and Arnaud (2001)
Pape (2010)
Bezzimyia yepezi Costa Rica, Puntarenas [NHMD] Pape and Arnaud (2001)
Bixinia collessi Australia [ANIC]
Bixinia winkleri Australia, Queenslad [ANIC]
Comoromyia sp. 1 Madagascar, Andringitra [MNHN]
Comoromyia sp. 2 Madagascar, Andringitra [MNHN]
Kinabalumyia pinax* Malaysia, Sabah [NHMUK]
Macrotarsina longimana* Croatia [MZUR]
Malayia fuscinervis Malaysia, Pahang and Philippines, Palawan [NHMD]
Marshallicona quitu* Ecuador [NHMD and MZUR]
Maurhinophora indoceanica* Mauritius [NHMUK]
Rhinophoridae Melanomyiodes capensis* South Africa, Western Cape [NHMUK]
Melanophora asetosa Israel [TAU] Pape (1986)
Cerretti and Pape (2009)
Melanophora basilewskyi Kenya [NHMD, TAU, MZUR] Crosskey (1977)
Pape (1986)
Cerretti and Pape (2009)
Melanophora chia Italy, Sardinia [MZUR] Cerretti and Pape (2009)
Melanophora roralis Italy: Latium and Veneto [MZUR] Crosskey (1977)
Pape (1986)
Pape (1998)
Cerretti and Pape (2009)
Metoplisa carbonaria Israel [MZUR and TAU]
Neotarsina andina* Peru [NHMD]
Neotarsina caraibica* Trinidad and Tobago [NHMD and NMHUK]
Oplisa tergestina Italia: Sicilia, Trentino-Alto Adige [MZUR] Tschorsnig (1985)
Pape (1986)
Parazamimus congolensis Burundi, Kayanza [NMB]
Paykullia insularis France, Corse [NHMD]
Paykullia maculata Czech Republic [NHMD] Tschorsnig (1985)
Pape (1986)
Phyto adolescens Italy, Sicilia [NHMD]
Phyto angustifrons Italy, Marche [TAU]
Queximyia flavipes South Africa: Eastern Cape and KwaZulu-Natal [NHMD] Crosskey (1977)
Pape (1986)
Rhinodonia antiqua New Caledonia [MNHN]
Rhinodonia flavicera New Caledonia [INHS]
Rhinomorinia sarcophagina Italy, Latium [MZUR] Tschorsnig (1985)
Pape (1986)
Rhinopeza gracilis New Guinea [ANIC]
Rhinophora lepida Italy, Veneto [MZUR]
Shannoniella cuspidata* Brazil, São Paulo [CNC] Nihei et al. (2016)
Shannoniella setinervis Brazil, São Paulo [CNC] Nihei et al. (2016)
Stevenia deceptoria* Italy, Sicily [MZUR]
Stevenia palermitana Italy, Sicily [MZUR] Cerretti and Pape (2007)
Tricogena rubricosa* Morocco [NHMD], Tunisia [NHMD]
Tromodesia angustifrons Israel [TAU], Greece [NHMD] Kugler (1978)
Pape (1986)
Trypetidomima fusca Nihei and Andrade (2014)
Trypetidomima lutea* Brasil, Rio de Janeiro [USNM]
Ventrops aethiopicus Ethiopia [TAU] Cerretti and Pape (2012)
Ventrops freidbergi Tanzania [TAU] Cerretti and Pape (2012)
Ventrops hannemariae Kenya [TAU], Malawi [TAU] Pape (1987)
Ventrops incisus Tanzania [NHMD] Pape (1987)
Ventrops intermedius Tanzania [TAU] Pape (1987)
Ventrops milichioides Tanzania and Kenya [TAU] Crosskey (1977)
Tschorsnig (1985)
Pape (1986)
Pape (1987)
Ventrops stuckenbergi Namibia and South Africa [MZUR] Cerretti and Pape (2012)

The data matrix was produced in Mesquite version 3.03 (Maddison and Maddison 2015). Inapplicable or unknown states were coded as ‘–’ and ‘?’, respectively (Table 3). Cladistic analyses were conducted with TNT version 1.5 (Goloboff and Catalano 2016). Heuristic searches were run under equal weights and under implied weighting (k-values: 3–10, 15, 20), with the ‘traditional search’ option under the following settings: General RAM of 1 GB, memory set to hold 1,000,000 trees, setting 1000 replicates with tree bisection-reconnection (TBR) branch swapping and saving 1000 trees per replicate. Multistate characters were treated as unordered. Character state changes (apomorphies) were optimised in WinClada version 1.00.08 (Nixon 2002) on the fittest tree obtained with a k-value of 4, using the unambiguous transformation algorithm. We chose the term “global apomorphies” for the uncontradicted and unreversed apomorphic character states, whereas “local apomorphies” was used for the homoplasious character states due to convergence or reversal.

Table 3.

Data matrix (outgroups in grey font).

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24
Musca spp. 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0
Amenia sp. 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0
Bengalia sp. 1 0 0 0 0 0 0 ? 0 ? ? 0 0 0 0 0 0 1 0 1 0 3
Calliphora vomitoria 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0
Melinda gentilis 1 0 0 0 0 0 0 0 0 1 0 0 0 ? 0 0 0 0 ? 2 0 0 0
Lucilia sericata 1 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0
Cuterebra austeni 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 2 4
Pollenia paupera 1 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 1 0
Eurychaeta muscaria 1 1 0 0 0 1 0 0 1 1 1 0 0 0 0 0 1 1 1 0 0 0 0 0
Rhyncomya impavida 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 1 1 0
Macquartia tenebricosa 0 1 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 1 0
Acompomintho lobata 0 0 0 0 1 0 0 0 1 1 0 0 0 0 0 0 1 1 0 0 0 0 2
Apomorphyto inbio 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0/1 0 0 0 0 0 2
Aporeomyia sp. 0 0 0 0 1 2 ? 0 0 ? 0 0 1 0 0 0 0 1 0 0 2
Axinia disjuncta 1 0 ? ? 0 ? 1 ? 0 0 0 0 1 0 0 0 0 ? 0 4
Axinia lucaris 0 0 0 0 1 0 0 0 1 1 0 0 0 1 1 0 1 0 0 0 0 0 2
Axinia miranda 0 0 0 0 ? 0 0 ? 2 ? 0 0 0 1 1 0 0 0 0 ? 0 3
Azaisia sp. 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 ? 0 0 0 0 2
Baniassa fascipennis 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 2
Bezzimyia barbarista 0 0 0 1 1 0 1 1 0 0 1 1 1 0 0 0 0 0 0 0 2
Bezzimyia bisecta 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 2
Bezzimyia busckii 0 0 0 0 1 0 1 1 0 0 1 1 1 0 0 0 0 0 0 0 2
Bezzimyia hansoni 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 2
Bezzimyia yepezi 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 0 0 0 2
Bixinia collessi 0 0 0 0 1 0 0 ? 0 0 0 0 0 1 0 0 0 0 0 0 2
Bixinia winckleri 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 2
Comoromyia sp. 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0/1 0 0 0 0 0 2
Comoromyia sp. 2 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 2
Kinabalumyia pinax 0 0 0 0 1 2 0 0 1 1 0 0 0 0 1 0 0 ? 0 1 0 2
Macrotarsina longimana 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 ? 0 0 0 0 2
Malayia fuscinervis 0 0 0 0 1 0 0 0 1 1 0 0 1 1 0 0 0/1 0 0/1 0 0 0 1 0
Marshallicona quitus 1 1 0 0 0 0 0 ? 1 ? 1 0 0 0 0 0 0 0 0 0 0 2
Maurhinophora indoceanica 0 0 0 0 0 0 0 0 ? 1 0 0 1 1 0 0 0 0 0 2 0 1 1
Melanomyiodes capensis 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 2
Melanophora asetosa 1 1 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 1 1 ? 0 2
Melanophora basilewskyi 1 1 0 0 0 0 0 ? 1 1 1 0 0 0 0 0 0 1 1 ? 0 3
Melanophora chia 1 1 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 1 0 0 0 2
Melanophora roralis 1 1 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 1 0 0 0 2
Metoplisa carbonaria 0 0 0 0 0 0 0 0 1 1 0/1 0 0 0 0 0 0 0 0 0 0 2
Neotarsina andina 0 0 0 0 0 0 0 1 0 0 ? 0 0 0 0 0 0 0 0 1 0 2
Neotarsina caraibica 0 0 0 0 0 0 0 1 0 0 ? 0 0 0 0 0 0 0 0 1 0 2
Oplisa tergestina 1 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 1 1
Parazamimus congolensis 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0 0 1 0 0 0 0 0 2
Paykullia insularis 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 2
Paykullia maculata 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 2
Phyto adolescens 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0
Phyto angustifrons 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 1 1 0 0 0 0 1 0
Queximyia flavipes 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 1
Rhinodonia antiqua 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 0 0 0 0 0 0 2
Rhinodonia flavicera 0 0 0 0 1 0 0 ? 1 ? 0 0 0 0 1 0 0 0 0 0 0 2
Rhinomorinia sarcophagina 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 1
Rhinopeza gracilis 0 0 0 0 1 0 0 ? 0 ? ? 0 0 0 0 0 0 0 1 0 0 3
Rhinophora lepida 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 1 0 1 0 0 0 2
Shannoniella cuspidata 0 0 0 0 1 1 0 0 2 2 1 1 1 1 0 0 0 0 0 0 0 2
Shannoniella setinervis 0 0 0 0 1 1 0 0 2 ? 1 1 1 1 0 0 0 0 0 0 0 2
Stevenia deceptoria 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 1 1 0 0 0 0 0 1
Stevenia palermitana 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 1 0 0 0 0 0 1
Tricogena rubricosa 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 1 1 0 0 0 0 0 1
Tromodesia angustifrons 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 2
Trypetidomima fusca 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2
Trypetidomima lutea 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2
Ventrops aethiopicus 0 0 0 0 0 0 0 ? 1 ? 1 0 0 0 0 0 1 0 0 0 0 0 1 0
Ventrops freidbergi 0 0 0 0 0 0 0 ? 1 ? 1 0 0 0 0 0 1 0 1 0 0 0 1 0
Ventrops hannemariae 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 1 0 0 0 0 0 2
Ventrops incisus 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 1 0 0 0 0 0 2
Ventrops intermedius 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 1 0 0 0 0 0 2
Ventrops milichioides 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0 1 0 0 0 0 0 2
Ventrops stuckenbergi 0 0 0 0 0 0 0 ? 1 ? 1 0 0 0 0 0 1 1 1 0 0 0 1 0
25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46
Musca spp. 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0
Amenia sp. 0 0 0 0 0 2 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0
Bengalia sp. 1 0 0 0 1 2 0 ? 0 0 0 0 0 ? 0 0 ? 0 0 0 0 0
Calliphora vomitoria 0 0 ? 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Melinda gentilis 0 0 0 0 1 2 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 0 0
Lucilia sericata 0 0 0 0 0 2 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 0 0
Cuterebra austeni 1 0 0 1 1 2 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0
Pollenia paupera 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Eurychaeta muscaria 0 0 0 0 1 2 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0
Rhyncomya impavida 0 0 ? 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Macquartia tenebricosa 0 2 0 0 1 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Acompomintho lobata 0 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Apomorphyto inbio 1 1 0 0 1 1 0 1 0 0 0 1 0 0 0 0 0 0 1 0 0
Aporeomyia sp. 1 1 0 0 1 0 0 ? 0 0 0 1 0 ? 0 0 ? 0 1 1
Axinia disjuncta 1 1 0 0 1 0 0 ? 0 1 0 1 0 ? 0 0 ? 0 1 1
Axinia lucaris 1 1 0 0 1 0 0 1 0 1 0 1 0 0 0 0 0 0 1 1
Axinia miranda 1 1 0 0 1 0 ? ? 0 1 0 1 0 ? 0 0 ? 0 1 1
Azaisia sp. 1 1 0 0 1 1 1 ? 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Baniassa fascipennis 0/1 1 0 0 0 2 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Bezzimyia barbarista 1 1 0 0 1 1 1 1 0 0 0 1 0 0 0 0 0 1 1
Bezzimyia bisecta 1 1 0 1 1 1 0 1 0 0 1 1 0 ? 0 0 ? 0 0 0
Bezzimyia busckii 1 1 0 0 1 1 1 1 0 0 0 1 0 0 0 0 0 1 1
Bezzimyia hansoni 1 1 0 1 1 1 0 ? 0 0 1 1 0 2 0 0 ? 0 0 0
Bezzimyia yepezi 1 1 0 0 1 2 0 0 0 0 1 1 0 0 0 0 0 0 0 0
Bixinia collessi 1 1 1 0 1 1 0 ? 0 1 1 1 0 0 0 0 0 0 1 0 1
Bixinia winckleri 1 1 1 0 1 1 1 1 0 1 1 1 0 0 0 0 0 0 1 0 1
Comoromyia sp. 1 2 1 0 1 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Comoromyia sp. 2 2 1 0 1 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Kinabalumyia pinax 1 1 0 0 1 0 0 ? 0 0 0 1 0 ? 0 0 ? 0 1 1
Macrotarsina longimana 1 1 0 0 1 1 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Malayia fuscinervis 1 1 0 0 1 1 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0
Marshallicona quitus 1 1 0 0 1 0 0 ? 0 0 0 1 0 ? 0 0 ? 2 1 0 0
Maurhinophora indoceanica 2 1 0 0 0 2 ? 0 0 0 0 1 0 0 0 ? 0 1 0 1 0 0
Melanomyiodes capensis 1 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Melanophora asetosa 1 1 0 1 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0
Melanophora basilewskyi 1 1 0 1 1 0 0 ? 0 1 0 1 0 ? 0 0 0 1 0 1
Melanophora chia 1 1 0 0 1 1 0 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0
Melanophora roralis 1 1 0 0 1 1 0 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0
Metoplisa carbonaria 1 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Neotarsina andina 1 1 0 0 1 0 1 1 1 0 0 1 0 0 0 0 0 2 1 0
Neotarsina caraibica 1 1 0 0 1 1 1 1 1 0 0 1 0 0 0 0 0 0 1 0
Oplisa tergestina 0 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Parazamimus congolensis 1 1 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Paykullia insularis 1 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Paykullia maculata 1 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Phyto adolescens 0 1 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Phyto angustifrons 0 1 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Queximyia flavipes 2 1 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Rhinodonia antiqua 0 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 1 1
Rhinodonia flavicera 0 1 0 0 1 1 0 ? 0 0 0 1 0 ? 0 0 ? 0 1 1
Rhinomorinia sarcophagina 0 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Rhinopeza gracilis 1 1 0 0 1 0 0 ? 0 1 0 1 0 ? 0 0 ? 0 0 0 1
Rhinophora lepida 1 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Shannoniella cuspidata 1 1 0 1 1 0 1 1 0 0 0 1 1 0 0 1 0 1 0 1 0 0
Shannoniella setinervis 1 1 0 1 1 0 1 ? 0 0 0 1 1 0 0 1 ? 1 0 1 0 0
Stevenia deceptoria 1 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Stevenia palermitana 0 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Tricogena rubricosa 1 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Tromodesia angustifrons 1 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Trypetidomima fusca 0 1 0 0 1 1 0 1 0 0 0 1 1 0 0 1 0 2 0 0 0 0
Trypetidomima lutea 1 1 0 0 1 1 0 1 0 0 0 1 1 0 0 1 0 2 0 0 0 0
Ventrops aethiopicus 1 1 0 0 1 1 0 0 0 0 0 1 0 ? 0 0 ? 0 0 0
Ventrops freidbergi 1 1 0 0 1 1 0 ? 0 0 0 1 0 ? 0 0 ? 0 0 0
Ventrops hannemariae 1 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Ventrops incisus 1 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 ? 0 0 0 0 0
Ventrops intermedius 1 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 ? 0 0 0 0 0
Ventrops milichioides 1 1 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
Ventrops stuckenbergi 1 1 0 0 1 1 0 0 0 0 0 1 0 ? 0 0 ? 0 0 0 0 0
47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68
Musca spp. 0 1 0 0 0 1 ? ? ? ? ? ? ? 0 0 0 1 0 1
Amenia sp. 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0
Bengalia sp. 0 1 0 0 1 1 1 1 0 0 0 0 0 1 0 1 0
Calliphora vomitoria 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0
Melinda gentilis 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0
Lucilia sericata 0 1 0 0 1 0 0 0 0 ? ? ? ? 0 0 0 0 1 0 1 0
Cuterebra austeni 0 1 0 0 1 1 1 1 0 1 0 0 0 1 0 0 0 0 0 1
Pollenia paupera 0 0 0 0 1 1 ? ? ? ? ? ? ? 0 0 0 0 0 0 1 0
Eurychaeta muscaria 0 1 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 0 1 1
Rhyncomya impavida 0 1 0 0 1 1 0 0 0 ? 3 0 0 0 0 0 0 0 0 0 1 0
Macquartia tenebricosa 0 1 0 0 0 0 ? ? ? ? ? ? ? 0 0 0 0 0 0 1 0
Acompomintho lobata 0 0 0 0 0 0 0 0 0 ? ? 0 ? 0 0 0 0 0 1 0 1 ?
Apomorphyto inbio 0 1 0 0 1 1 0 1 0 0 0 0 0 0 0 0 0 0 1 1 1 0
Aporeomyia sp. 0 1 0 1 0 0 0 1 0 0 0 1 0 0 1 0 0 1 0 0 1 0
Axinia disjuncta 0 1 0 0 1 1 0 0 0 0 0 0 0 0 1 1 0 0 1 1 0 1
Axinia lucaris 0 1 0 0 1 1 0 0 1 0 0 0 1 0 1 1 0 0 0 1 1 1
Axinia miranda 0 1 0 0 1 1 0 0 0 0 0 0 1 0 1 1 0 0 1 1 1 ?
Azaisia sp. 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1
Baniassa fascipennis 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0
Bezzimyia barbarista 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0
Bezzimyia bisecta 1 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1 1 0
Bezzimyia busckii 1 0 0 1 1 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 0
Bezzimyia hansoni 1 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1 1 0
Bezzimyia yepezi 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 1 1 0
Bixinia collessi 0 1 0 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 1 0 1 0
Bixinia winckleri 0 1 0 1 1 1 0 0 0 1 0 0 0 0 0 0 0 0 1 0 1 0
Comoromyia sp. 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0
Comoromyia sp. 2 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0
Kinabalumyia pinax 0 1 0 1 0 0 1 0 0 0 0 1 1 0 1 0 0 0 0 0 1 0
Macrotarsina longimana 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1
Malayia fuscinervis 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0
Marshallicona quitus 0 1 1 0 1 1 0 0 0 1 2 0 0 0 0 0 0 0 1 0 1 1
Maurhinophora indoceanica 0 1 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?
Melanomyiodes capensis 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0
Melanophora asetosa 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0
Melanophora basilewskyi 1 0 0 1 1 0 0 0 0 0 0 0 0 1 0 0 1 0 1 1 0
Melanophora chia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0
Melanophora roralis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0
Metoplisa carbonaria 0 1 0 0 1 0 0 0 0 0 2 0 0 0 0 0 0 0 1 0 0 0
Neotarsina andina 0 1 0 0 1 1 0 0 0 0 0 0 1 0 0 0 0 0 0 1 1 0
Neotarsina caraibica 0 1 0 0 1 1 0 0 0 0 1 0 1 0 0 0 0 0 0 1 1 0
Oplisa tergestina 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1
Parazamimus congolensis 0 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0
Paykullia insularis 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0
Paykullia maculata 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0
Phyto adolescens 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0
Phyto angustifrons 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0
Queximyia flavipes 0 1 0 0 0 0 0 0 0 ? 3 0 0 0 0 0 0 0 1 0 0 0
Rhinodonia antiqua 0 1 0 0 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 1 0
Rhinodonia flavicera 0 1 0 0 1 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 1 0
Rhinomorinia sarcophagina 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0
Rhinopeza gracilis 0 1 0 0 0 0 0 0 0 ? ? ? ? 0 0 0 0 0 0 1 1 0
Rhinophora lepida 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 1 0
Shannoniella cuspidata 0 1 1 0 1 1 0 0 0 0 2 0 0 0 0 0 0 0 1 1 1 0
Shannoniella setinervis 0 1 1 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 1 1 1 0
Stevenia deceptoria 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1
Stevenia palermitana 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1
Tricogena rubricosa 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1
Tromodesia angustifrons 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0
Trypetidomima fusca 0 1 1 0 0 0 0 0 0 ? 3 ? 0 0 0 0 0 0 1 1 1 ?
Trypetidomima lutea 0 1 1 0 0 0 0 0 0 ? 3 ? 0 0 0 0 0 0 1 1 1 ?
Ventrops aethiopicus 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0
Ventrops freidbergi 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0
Ventrops hannemariae 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 1 0
Ventrops incisus 0 1 0 0 0 0 0 0 0 1 2 0 0 0 0 0 0 0 1 0 1 0
Ventrops intermedius 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 1 0
Ventrops milichioides 0 1 0 0 0 0 0 0 0 1 2 0 0 0 0 0 0 0 1 0 1 0
Ventrops stuckenbergi 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 1 0
69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91
Musca spp. 1 ? 0 ? 0 0 0 0 0 0 1
Amenia sp. 0 0 0 0 0 1 0 0 0 ? ? ? ? ? ? ? ? ? ?
Bengalia sp. 0 0 0 1 0 1 ? 0 ? 0 0 0 0 0 0 1
Calliphora vomitoria 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Melinda gentilis 0 0 0 0 0 1 0 0 0 0 1 ? 0 0 0 0 1 0 0
Lucilia sericata 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 ? 0 0 0 0 0 0 1
Cuterebra austeni 0 0 0 0 1 0 0 0 ? ? ? ? ? ? ? ? ? ?
Pollenia paupera 0 0 0 0 0 0 1 0 0 0 0 0 0 ? ? ? 0 0 0 0 0 0 0
Eurychaeta muscaria 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? 0 ? ?
Rhyncomya impavida 0 0 1 0 0 0 0 0 0 0 0 0 0 0 ? 0 ? ? ? ? ? ? ?
Macquartia tenebricosa ? ? ? ? ? ? ? ? ? ?
Acompomintho lobata 0 0 0 0 0 0 1 0 0 0 1 0 0 0 ? ? ? ? ? ? ? ? ?
Apomorphyto inbio 0 0 0 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ?
Aporeomyia sp. 1 0 0 0 0 1 0 0 0 ? ? ? ? ? ? ? ? ? ?
Axinia disjuncta 1 0 1 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ?
Axinia lucaris 1 0 1 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ?
Axinia miranda ? 1 ? ? 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ?
Azaisia sp. 0 0 0 0 1 0 0 0 0 0 1 0 ? 0 ? ? ? ? ? ? ? ? ?
Baniassa fascipennis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ?
Bezzimyia barbarista 0 0 0 0 0 1 ? ? ? ? 0 0 0 1 ? ? ? ? ? ? ? ? ?
Bezzimyia bisecta 1 0 1 0 0 1 ? ? ? ? 0 0 0 0 ? ? ? ? ? ? ? ? ?
Bezzimyia busckii 0 0 0 0 0 0 1 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ?
Bezzimyia hansoni 1 0 1 0 0 1 ? ? ? ? 0 0 0 0 ? ? ? ? ? ? ? ? ?
Bezzimyia yepezi 1 0 1 0 0 1 ? ? ? ? 0 0 0 1 1 1 1 1 0 1 ? 0 1
Bixinia collessi 1 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ?
Bixinia winckleri 1 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ?
Comoromyia sp. 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ?
Comoromyia sp. 2 0 0 0 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ?
Kinabalumyia pinax 1 0 0 0 0 0 1 1 0 0 0 0 1 1 ? ? ? ? ? ? ? ? ?
Macrotarsina longimana 0 0 0 0 0 0 0 0 0 0 1 0 0 0 ? ? ? ? ? ? ? ? ?
Malayia fuscinervis 0 0 0 0 0 1 ? ? ? ? 0 0 0 1 ? ? ? ? ? ? ? ? ?
Marshallicona quitus 1 0 1 0 0 0 0 1 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ?
Maurhinophora indoceanica ? ? ? ? ? ? ? ? ? ? ? ? 0 0 ? ? ? ? ? ? ? ? ?
Melanomyiodes capensis ? 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ?
Melanophora asetosa 0 0 1 0 0 0 0 0 0 0 0 0 0 1 ? ? ? ? ? ? ? ? ?
Melanophora basilewskyi 1 1 ? ? 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ?
Melanophora chia 0 0 1 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ?
Melanophora roralis 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1 2 1 1 0 1 ? 0 1
Metoplisa carbonaria 0 0 0 0/1 1 0 1 0 0 0 1 0 0 0 ? ? ? ? ? ? ? ? ?
Neotarsina andina 1 0 0 1 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ?
Neotarsina caraibica 1 0 0 1 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ?
Oplisa tergestina 0 0 0 0 1 0 0 0 0 0 1 0 0 0 1 1 1 1 0 1 ? 1 1
Parazamimus congolensis 1 0 0 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ?
Paykullia insularis 1 0 0 1 0 0 0 0 0 0 0 0 0 1 1 2 1 1 0 1 ? 0 1
Paykullia maculata 0 0 0 1 0 0 0 0 0 0 0 0 0 1 1 2 1 1 0 1 ? 0 1
Phyto adolescens 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 2 1 1 0 1 ? 0 1
Phyto angustifrons 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 2 1 1 0 1 ? 0 1
Queximyia flavipes 0 0 1 0 0 0 0 0 0 0 1 0 0 0 ? ? ? ? ? ? ? ? ?
Rhinodonia antiqua 0 0 1 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ?
Rhinodonia flavicera 0 0 1 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ?
Rhinomorinia sarcophagina 0 0 0 0 1 0 0 0 0 0 1 0 0 0 1 1 1 1 0 1 ? 1 1
Rhinopeza gracilis 0 0 0 0 0 0 1 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ?
Rhinophora lepida 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 1 ? 1 1
Shannoniella cuspidata 0 0 1 0 0 0 0 1 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ?
Shannoniella setinervis 0 0 1 0 0 0 0 1 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ?
Stevenia deceptoria 0 0 0 0 1 0 1 0 0 0 1 0 0 0 1 1 1 1 0 1 ? 1 1
Stevenia palermitana 0 0 0 0 1 0 1 0 0 0 1 0 0 0 1 1 1 1 0 1 ? 1 1
Tricogena rubricosa 0 0 0 0 1 0 1 0 0 0 1 0 0 0 1 1 1 1 0 1 ? 1 1
Tromodesia angustifrons 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ?
Trypetidomima fusca 0 0 1 0 0 0 0 1 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ?
Trypetidomima lutea 0 0 1 0 0 0 0 1 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ?
Ventrops aethiopicus 0 0 1 0 0 0 0 0 0 0 0 1 0 ? ? ? ? ? ? ? ? ? ?
Ventrops freidbergi 0 0 1 0 0 0 0 0 0 0 0 1 0 ? ? ? ? ? ? ? ? ? ?
Ventrops hannemariae 0 0 1 0 0 0 0 1 1 1 0 1 0 0 ? ? ? ? ? ? ? ? ?
Ventrops incisus 0 0 1 0 0 0 0 0 0 0 0 1 0 0 ? ? ? ? ? ? ? ? ?
Ventrops intermedius 0 0 1 0 0 0 0 1 1 1 0 1 0 0 ? ? ? ? ? ? ? ? ?
Ventrops milichioides 0 0 1 0 0 0 0 0 0 0 0 1 0 0 ? ? ? ? ? ? ? ? ?
Ventrops stuckenbergi 0 0 1 0 0 0 0 1 1 0 0 1 0 ? ? ? ? ? ? ? ? ? ?
92 93 94 95 96 97 98 99 92 93 94 95 96 97 98 99
Musca spp. 0 0 0 0 0 1 0 Melanophora asetosa ? ? ? ? ? ? ? ?
Amenia sp. ? ? ? ? ? ? ? ? Melanophora basilewskyi ? ? ? ? ? ? ? ?
Bengalia sp. 0 0 0 ? 0 1 0 Melanophora chia ? ? ? ? ? ? ? ?
Calliphora vomitoria 0 0 0 0 0 0 ? ? Melanophora roralis 1 0 0 1 1 0 0 1
Melinda gentilis 1 0 0 ? 0 ? ? 1 Metoplisa carbonaria ? ? ? ? ? ? ? ?
Lucilia sericata 0 0 0 ? 0 0 Neotarsina andina ? ? ? ? ? ? ? ?
Cuterebra austeni ? ? ? ? ? ? ? ? Neotarsina caraibica ? ? ? ? ? ? ? ?
Pollenia paupera 0 0 0 ? 0 ? ? Oplisa tergestina 0 1 0 0 0 1 0 0
Eurychaeta muscaria ? ? ? ? ? ? ? ? Parazamimus congolensis ? ? ? ? ? ? ? ?
Rhyncomya impavida ? ? ? ? ? ? ? ? Paykullia insularis 1 0 0 1 1 0 0 1
Macquartia tenebricosa ? ? ? ? ? ? ? ? Paykullia maculata 1 0 0 1 1 0 0 1
Acompomintho lobata ? ? ? ? ? ? ? ? Phyto adolescens 1 0 0 1 1 0 0 1
Apomorphyto inbio ? ? ? ? ? ? ? ? Phyto angustifrons 1 0 0 1 1 0 0 1
Aporeomyia sp. ? ? ? ? ? ? ? ? Queximyia flavipes ? ? ? ? ? ? ? ?
Axinia disjuncta ? ? ? ? ? ? ? ? Rhinodonia antiqua ? ? ? ? ? ? ? ?
Axinia lucaris ? ? ? ? ? ? ? 1 Rhinodonia flavicera ? ? ? ? ? ? ? ?
Axinia miranda ? ? ? ? ? ? ? ? Rhinomorinia sarcophagina 0 1 0 0 0 1 0 0
Azaisia sp. ? ? ? ? ? ? ? ? Rhinopeza gracilis ? ? ? ? ? ? ? ?
Baniassa fascipennis ? ? ? ? ? ? ? ? Rhinophora lepida 0 1 0 0 0 1 0 0
Bezzimyia barbarista ? ? ? ? ? ? ? ? Shannoniella cuspidata ? ? ? ? ? ? ? ?
Bezzimyia bisecta ? ? ? ? ? ? ? ? Shannoniella setinervis ? ? ? ? ? ? ? ?
Bezzimyia busckii ? ? ? ? ? ? ? ? Stevenia deceptoria 0 1 0 0 0 1 0 0
Bezzimyia hansoni ? ? ? ? ? ? ? ? Stevenia palermitana 0 1 0 0 0 1 0 0
Bezzimyia yepezi 0 1 1 0 0 1 1 0 Tricogena rubricosa 0 1 0 0 0 1 0 0
Bixinia collessi ? ? ? ? ? ? ? ? Tromodesia angustifrons ? ? ? ? ? ? ? ?
Bixinia winckleri ? ? ? ? ? ? ? ? Trypetidomima fusca ? ? ? ? ? ? ? ?
Comoromyia sp. 1 ? ? ? ? ? ? ? ? Trypetidomima lutea ? ? ? ? ? ? ? ?
Comoromyia sp. 2 ? ? ? ? ? ? ? ? Ventrops aethiopicus ? ? ? ? ? ? ? ?
Kinabalumyia pinax ? ? ? ? ? ? ? ? Ventrops freidbergi ? ? ? ? ? ? ? ?
Macrotarsina longimana ? ? ? ? ? ? ? ? Ventrops hannemariae ? ? ? ? ? ? ? ?
Malayia fuscinervis ? ? ? ? ? ? ? ? Ventrops incisus ? ? ? ? ? ? ? ?
Marshallicona quitus ? ? ? ? ? ? ? ? Ventrops intermedius ? ? ? ? ? ? ? ?
Maurhinophora indoceanica ? ? ? ? ? ? ? ? Ventrops milichioides ? ? ? ? ? ? ? ?
Melanomyiodes capensis ? ? ? ? ? ? ? ? Ventrops stuckenbergi ? ? ? ? ? ? ? ?

Catalogue

Format

The present catalogue lists all nominal genera and species of Rhinophoridae, providing details about name-bearing types and with known distributions updated from both recent literature and our own identifications of museum specimens.

Valid taxa are arranged hierarchically and alphabetically, according to genus and species (subfamilial and tribal classification is considered premature given the difficulties in interpreting adult homologies and defining monophyletic groupings). Synonyms, including unjustified emendations and incorrect original and subsequent spellings, are listed chronologically for all names.

Each genus-group name is listed with the following formatting and information: genus name (in square brackets if unavailable, italics if available, bold + italics if valid), author, year, page, type species with author and date, form of type fixation with author and date. Each type species is given in its original binomen (Recommendation 67B of the “International Code of Zoological Nomenclature”, henceforth “the Code”, ICZN 1999), followed by its valid name, if different, in square brackets. Incorrect original spellings are given teste their First Reviser. Incorrect subsequent spellings encountered during this study are cited from their earliest occurrence.

Species are listed alphabetically by valid name followed by synonyms, nomina nuda, unjustified emendations and incorrect spellings listed chronologically. The genus Bezzimyia Townsend is likely polyphyletic and the species have been grouped into two species groups (namely Group A and Group B) and listed alphabetically within each group. The valid specific epithet is given in bold and italics followed by author and year. Each available name is given in italics in its original combination and spelling followed by author, year (with letter if applicable, to match with References), and page. Given next is the type locality in modern spelling, followed by information about the name-bearing type, consisting of status (holotype, lectotype, neotype or syntypes), sex, and acronym of type repository. Additional information may be given under “Remarks”. Distribution is given hierarchically and alphabetically according to biogeographical region and by country, but with larger countries separated into states/provinces and offshore islands listed separately from the mainland. Archipelagos may be listed by island when data are available. European distribution follows Fauna Europaea (https://fauna-eu.org/, see Pape et al. 2015).

Type localities are cited from largest to smallest geographic area or place. Country and state/province names are given only in their modern equivalents. Coordinates given in an original publication are cited as an integral part of the type locality, in their original format.

For data on the number and sex of name-bearing types other than an unambiguous fixation of a holo-, lecto- or neotype, we follow the format proposed by O’Hara and Cerretti (2016), slightly modified as explained below.

Type(s), male: One or more males. This citation is used for a species described from the male sex without indication whether the type series comprised a single male (i.e., a holotype) or more than one male (i.e., syntypes).

Type(s), female: One or more females.

Type(s), unspecified sex: One or more specimens with no indication of sex.

Syntypes, [number] male[s] and [number] female[s] (e.g., “3 males and 2 females”): Species described from both sexes, with the exact number of males and females specified and without a designated holotype.

Syntypes, males and females: Species described from both sexes, with more than one specimen of each sex but without specified numbers and without a designated holotype.

Syntypes, male(s) and female(s): Species described from both sexes, with no indication of the number of specimens of either sex, neither the exact number nor whether only one or more than one.

Syntypes, males: Species described from more than one male, without indication of the specific number of males and without a designated holotype.

Syntypes, females: Species described from more than one female, without indication of the specific number of females and without a designated holotype.

Syntypes, unspecified number and sex: Species described from more than one specimen but without indication of sex or number of specimens and without a designated holotype.

Avoidance of assumption of holotype and lectotypifications

Recommendation 73F of the Code (ICZN 1999), “Avoidance of assumption of holotype”, recommends, “where appropriate” and when it is possible that the nominal species-group taxon was based on more than one specimen, to designate a lectotype rather than assume a holotype; and Article 74.6 of the Code deems that an assumed monotypy where the original description neither implies nor requires that there were syntypes, is deemed to be a lectotype designation if it is considered subsequently that the original description was based on more than one specimen. We follow O’Hara et al. (2009) and O’Hara and Cerretti (2016) in using the term “lectotype designation” for an explicit lectotypification where the author used either the term “lectotype” or an exact translation or equivalent expression (e.g., “the type”), and the term “lectotype fixation” for an implicit lectotypification by inference of holotype as well as for cases where the original work reveals that the taxon had been based on more than one specimen none of which were designated as holotype and an author subsequently used the term “holotype” in a way that explicitly indicated that he or she was selecting from the type series that particular specimen to serve as the name-bearing type.

Distributional data

Distributions are cited for each valid species based on published records, examination of specimens in collections, and material collected by the authors or made available by colleagues. New country records are followed by “[new record]”, and the label data of the relevant specimen(s) upon which the new record is based are given in Table 4. The primary sources for distributions were Cerretti (2002, 2003, 2007), Colless (1994), Crosskey (1977), Draber-Mońko (2007), Ebejer (2011), Ginn (2012), Guimarães (1971), Herting (1961, 1993), Kugler (1978), Martínez and Nihei (2018), Mulieri et al. (2010), O’Hara et al. (2015), Pape (1998a, 1998b, 2010), Peris (1963), Rognes and Hansen (1996), Sabrosky and Arnaud (1965), Verves (2005, 2010, 2012), Verves and Khrokalo (2006, 2010), Verves et al. (2019), Wood (1987), and Zeegers (2008, 2011).

Table 4.

New country records.

Species Locality Source / repository
Acompomintho sinensis Tajikistan, Gorno-Badachšan, Rŭshan NHMD
Macrotarsina longimana Italy, Sicily, Palermo province, Bisacquino, Riserva Naturale Monte Genuardo 900 m, 12.VIII.2000, P. Cerretti leg. MZUR
Melanophora roralis Canada, Ontario http://bugguide.net/node/view/443145
USA, Ohio http://www.inaturalist.org/observations/101858
British Virgin Islands, St Thomas, Charlotte Amalie AMNH
Oplisa aterrima Italy, Sicily, Palermo province, Bosco della Ficuzza 600–1000 m, Torretta Torre, 18.V.2004, P. Cerretti leg. MZUR
Portugal, Coimbra, Buçaco Forest, 5.vii.1990, V. Michelsen (NHMD) NHMD
Paykullia braueri Croatia, Ličko-senjaska Co., 4 km NW Rudelić Draga, 280m, 44°28'41.92"N, 15°8'57.34"E, 14.vi.2012, E. Buenaventura, T. Pape, D. Whitmore NHMD
Paykullia maculata Spain, Granada, Trevélez (S Mulhacén), 1480m, 9.vii.1993, V. Michelsen NHMD
Phyto abbreviata Tunisia, 25 km E Gafsa, 11–13.iii.1986, Zool. Mus. Copenhagen Exp. NHMD
Phyto adolescens Greece, Pelopónnisos, Taïyetos Mts, 950–1800 m, 15–19.v.1990, Zool. Mus. Copenhagen Exp. NHMD
Phyto discrepans Malta, Buskett Garden, Rabat, 4–11.vi.1988, Stig Andersen NHMD
Morocco, 600 m, Checheouèn, 22.iv.1989, Zool. Mus. Copenhagen Exp. NHMD
Morocco, 300 m, Ouezzane, 21–22.iv.1989, Zool. Mus. Copenhagen Exp. NHMD
Morocco, 1150 m, 40 km N Fès, 20.iv.1989, Zool. Mus. Copenhagen Exp. NHMD
Portugal, Coimbra, Buçaco Forest, 5.vii.1990, V. Michelsen NHMD
Tunisia, 25 km SE Ain Draham, 10–16.v.1988, Zool. Mus. Copenhagen Exp. NHMD
Tunisia, Tabarka area, 7–18.v.1998, Zool. Mus. Copenhagen Exp. NHMD
Tunisia, 40 km, W Jendouba 17.v.1988, Zool. Mus. Copenhagen Exp. NHMD
Phyto discrepans Tunisia, El Kef area, 14.v.1988, Zool. Mus. Copenhagen Exp. NHMD
Tunisia, 15 km NW Kebili, 17.iii.1986, Zool. Mus. Copenhagen Exp. NHMD
Phyto melanocephala Croatia, Ličko-senjaska Co., 2.8 km SSE Sveti Juraj, 380m, 44°54'28.38"N, 14°56'26.84"E, 12.vi.2012, E. Buenaventura, T. Pape, D. Whitmore NHMD
Croatia, Ličko-senjaska Co., nr. Sušanj Cesarički, 850 m, 44°31'51.39"N, 15°7'37.46"E, 13.vi.2012, E. Buenaventura, T. Pape, D. Whitmore NHMD
Greece, Makedonia/Tessalia, Olympos 700–2100 m, 21–26.v.1990, Zool. Mus. Copenhagen Exp. NHMD
Sweden, Möckelmossen, Vickelby, 1.vi.2006, V. Michelsen NHMD
Turkey, Pamphylia, W of Alanya, 2–13.vi.1991, B. Petersen leg. NHMD
Rhinophora lepida Italy, Trentino-Alto Adige, Trento prov., Strada da Sdruzzinà a Villaggio san Michele, 810 m, 45°43'31"N, 10°58'01"W, 26–28.VIII.2015, D. Corcos MZUR
Italy, Veneto, Belluno prov., Misurina, 1400 m, 46°32'54.08"N, 12°14'51.08"W, 30.VIII–1.IX.2015, D. Corcos MZUR
Italy, Veneto, Belluno prov., Misurina, 1500 m, 46°33'21.26"N, 12°14'48.26"W, 30.VII–1.VIII.2015, D. Corcos MZUR
Italy, Veneto, Belluno prov., Misurina, 1600 m, 46°33'43.96"N, 12°14'41.61"W, 30.VII–1.VIII.2015, D. Corcos MZUR
Italy, Veneto, Belluno prov., Cortina-Passo Falzarego, 1300 m, 46°32'7.30"N, 12°7'25.25"W, 30.VII–1.VIII.2015, D. Corcos MZUR
Italy, Veneto, Belluno prov., Cortina-Passo Falzarego, 2100 m, 46°31'8.67"N, 12°0'40.55"W, 30.VII–1.VIII.2015, D. Corcos MZUR
Italy, Veneto, Belluno prov., Cortina-Passo Giau, 1800 m, 46°29'51.45"N, 12°4'33.87"W, 30.VII–1.VIII.2015, D. Corcos MZUR
Italy, Veneto, Belluno prov., Cortina-Passo Giau, 1900 m, 46°29'32.44"N, 12°4'20.13"W, 30.VII–1.VIII.2015, D. Corcos MZUR
Stevenia atramentaria Greece, Pelopónnisos, Taïyetos Mts, 950–1800 m, 15–19.v.1990, Zool. Mus. Copenhagen Exp. NHMD
Stevenia bertei Croatia, Ličko-senjaska Co., nr. Sušanj Cesarički, 850 m, 44°31'51.39"N, 15°7'37.46"E, 13.vi.2012, E. Buenaventura, T. Pape, D. Whitmore NHMD
Croatia, Ličko-senjaska Co., nr. Baške Oštarije, 920m, 44°31'34.50"N, 15°10'5.50"E, 13.vi.2012, E. Buenaventura, T. Pape, D. Whitmore NHMD
Croatia, Ličko-senjaska Co., nr. Podoštra, 665m, 44°31'48.63"N, 15°19'35.92"E, 15.vi.2012, E. Buenaventura, T. Pape, D. Whitmore NHMD
Stevenia hirtigena Israel, Negev, Ein Avdat NP, 25.v.2004, K. Szpila NHMD
Stevenia signata Turkey, Pamphylia, W of Alanya, 2–13.vi.1991, B. Petersen leg. NHMD
Stevenia umbratica Tunisia, Ain Drahanm area, 5–18.v.1988, Zool. Mus. Copenhagen Exp. NHMD
Tromodesia setiventris Pakistan, Normal (nr. Skardu), NWFP. 1988.8.16, T. Hayashi NHMD

Collections

AMNH American Museum of Natural History, New York, USA.

ANIC Australian National Insect Collection, CSIRO, Canberra, Australia.

NBCL Naturalis Biodiversity Center, Leiden (including collections formerly deposited at Zoölogisch Museum, Universiteit van Amsterdam), Netherlands.

BLKU Biosystematics Laboratory, Kyushu University, Fukuoka, Japan.

BPBM Bernice Pauahi Bishop Museum, Honolulu, Hawaii, USA.

CNC Canadian National Collection of Insects, Agriculture and Agri-Food Canada, Ottawa, Canada.

CULSP Czech University of Life Sciences, Prague, Czech Republic.

DEI Deutsches Entomologisches Institut, Leibniz-Zentrum für Agrarlandschaftsforschung, Müncheberg, Germany.

ENSAM École nationale supérieure agronomique de Montpellier, France.

INBio Instituto Nacional de Biodiversidad, San José, Costa Rica [INBio collections are now under the care of the Museu Nacional de Costa Rica, San José].

IRSNB Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium.

LSUK Linnean Society, London, United Kingdom.

MIZA Museo del Instituto de Zoología Agrícola, Maracay, Venezuela.

MNHN Muséum national d’Histoire naturelle, Paris, France.

MNCN Museo Nacional de Ciencias Naturales, Madrid, Spain.

MRAC Musée Royal de l’Afrique Centrale.

MZF Museo Zoologico “La Specola”, Florence, Italy.

MZLU Museum of Zoology, Lund University, Lund, Sweden.

MZSP Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil.

MZUR Museo di Zoologia. Sapienza Università di Roma, Rome, Italy.

NHMD Natural History Museum of Denmark, University of Copenhagen, Copenhagen, Denmark. [Formerly as ZMUC.]

NHMUK Natural History Museum [formerly British Museum (Natural History)], London, United Kingdom.

NHMW Naturhistorisches Museum Wien, Vienna, Austria.

NHRS Naturhistoriska riksmuseet, Stockholm, Sweden.

NMBA Naturhistorisches Museum der Benediktiner-Abtei Admont, Admont, Austria.

NMDA Department of Arthropoda, KwaZulu-Natal Museum, Pietermaritzburg, South Africa.

NMNW National Museum of Namibia, Windhoek, Namibia.

PUCE Pontificia Universidad Cátolica Ecuador, Quito, Ecuador.

QDPC Queensland Department of Primary Industries, Indooroopilly, Queensland, Australia.

SAMC Iziko South African Museum, Cape Town, South Africa.

SMNS Staatliches Museum für Naturkunde Stuttgart, Stuttgart, Germany.

TAU The Steinhardt Museum of Natural History, Tel Aviv University, Tel Aviv, Israel.

TFMC Museo de Ciencias Naturales de Santa Cruz de Tenerife, Canary Islands, Spain.

UASK National Academy of Sciences of Ukraine, Kiev, Ukraine.

USNM National Museum of Natural History [formerly United States National Museum], Smithsonian Institution, Washington DC, USA.

ZIN Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia.

ZMHB Museum of Natural History, Leibniz-Institute for Research on Evolution and Biodiversity, Berlin, Germany.

ZMUM Zoological Museum of Moscow University, Moscow, Russia.

Key to world genera of Rhinophoridae

The following is a standard dichotomous key with couplets containing the main characters chosen for the key path, with alternative states in the respective entries of the couplet, listed by their supposed diagnostic strength. Occasionally, supplementary information is given in square brackets where considered helpful to secure an identification. The key includes the genus Alvamaja as well as the Afrotropical Moriniacarinata species group” (Pape 1997, Rognes 2010), all of which were recently transferred from Rhinophoridae to the re-established oestroid family Polleniidae (Cerretti et al. 2019). As these polleniids are nearly indistinguishable from rhinophorids, and given the lack of comprehensive keys to world oestroid families and genera, their inclusion is deemed helpful. They are given within square brackets. The key also includes the genus Aporeomyia, which was originally tentatively assigned to the Tachinidae based on the morphology of the phallus, but which is here moved to Rhinophoridae based on a re-assessment of the homology of relevant phallic sclerotisations (see below).

1 Body ground colour metallic green or blue-violet. Postalar wall with a tuft of fine setulae [Alvamaja Rognes; Polleniidae]
Body ground colour not metallic green or blue-violet, at most shiny black. Postalar wall bare [if with short setae then not Rhinophoridae, see couplets 8, 13, 17, 22] 2
2 Brachypterous, micropterous or apterous specimen (Fig. 4H), female only 3
Wing fully developed, male or female 4
3 Tergite 7 and sternite 7 forming a sclerotised, dorsoventrally flattened oviscapt (Fig. 13K–N, or similar structures) Axinia Colless (in part)
Segment 7 normally developed, i.e., not modified into a dorsoventrally flattened oviscapt Bezzimyia Townsend (in part)
4 Vein M1 running nearly straight or evenly curved forward to wing margin (Figs 9B, C; 10D, M, N; 16I; 17J; 18F), i.e., without a distinct bend 5
Vein M1 with a distinct, sometimes shallow bend that can be well removed from wing margin (e.g., Figs 9A, D–F, K, L; 10B, C, E–L) or very close to it (Fig. 9G), or M1 not reaching wing margin (i.e., ending freely in the wing membrane) (Fig. 9H–J) 11
5 Shiny black or brown flies, usually with bright yellow antenna or, more rarely, with variously patterned legs; with very short setae on head, scutum and abdomen, of approx. the same size as the smallest clothing setae (Fig. 4C–E). Vein CuA+CuP not reaching wing margin. Male: postpedicel characteristically rounded-subtriangular (axe-head-like and shorter than maximum distal width) (Fig. 6C–F), the thickened posterior margin abutting the enlarged, sunken face or anterior margin forming two or more lobes; arista very short (i.e., distinctly shorter than length of postpedicel) to nearly absent, and arising apically or sub-apically on postpedicel (Fig. 6C–F). Female: tergite 7 and sternite 7 forming a strongly sclerotised, dorsoventrally flattened and evenly curved, usually long oviscapt (Fig. 13K–N) Axinia Colless
Body colour varies from black to yellow, microtomentum present or absent; setae of head and thorax usually normally developed. Male: postpedicel usually not triangular in lateral view (if so, then postpedicel much longer than its maximum distal width and vein CuA+CuP reaching wing margin), arista arising in proximal half (Fig. 18C, E). Female: oviscapt not modified 6
6 Antenna short, at most as long as minimum height of gena (usually much shorter) (Figs 16D, F; 17F, H). Antenna arising distinctly below middle of compound eye. Tarsus of fore leg strongly laterally compressed (Figs 16A, H; 17B, E). Palpus strongly reduced, approx. 1–2 times as long as wide Neotarsina Cerretti & Pape, gen. nov.
Antenna much longer than minimum height of gena, arising at or above middle of compound eye. Tarsus of fore leg not laterally compressed. Palpus usually well developed, if reduced then mouthparts vestigial 7
7 First postsutural supra-alar seta well developed, at least as long and robust as posterior notopleural seta (Fig. 13B, F) 8
First postsutural supra-alar seta absent or minute (Fig. 13E) 9
8 Head longer than high and facial profile not receding. Prementum at least as long as head height and labella narrow. Antenna brown to black. Upper half of facial plate carinate (Fig. 13A). Postalar wall usually with at least one small setula. Male: cerci normally developed and medially not fused into a syncercus; surstylus articulated (i.e., not fused) to epandrium; extension of dorsal sclerite of distiphallus divided medially into two sclerites; median process of ventral sclerotisation of distiphallus not fused to base of ventral sclerotisation [Moriniacarinata species group” (in part); Polleniidae]
Head higher than long (even if slightly so) and facial profile receding (Figs 5E; 7H). Prementum usually shorter than height of head, if approx. as long as head height and labella narrow, then antenna pale yellow. Face not carinate. Postalar wall bare. Male: cerci very short and medially fused into a syncercus; surstylus fused to epandrium; extension of dorsal sclerite of distiphallus not divided medially; median process of ventral sclerotisation of distiphallus very long, narrow and fused to base of ventral sclerotisation Rhinodonia Cerretti, Lo Giudice & Pape
9 Vein CuA+CuP not reaching wing margin (Fig. 10M). First aristomere at most as long as wide, second aristomere approx. 4 times as long as wide. Mouthparts strongly reduced (vestigial). Male: proclinate orbital setae absent; postpedicel normal, i.e., not divided longitudinally into lobes; cerci not fused medially, distally well divided into two pointed branches; surstylus fused to epandrium Rhinopeza Cerretti, Lo Giudice & Pape
Vein CuA+CuP reaching wing margin (Fig. 9B). First aristomere at least 4 times as long as wide, second aristomere at least 8 times as long as wide (at least half as long as third aristomere). Mouthparts normally developed or strongly reduced. Male: 0–1 proclinate orbital setae; postpedicel varying from normal to branching into three lobes; cerci medially fused into a syncercus or distally well-divided into two pointed branches; surstylus not fused to epandrium 10
10 Mouthparts normally developed (Fig. 18A, C, E). Male: 1 proclinate orbital seta; postpedicel whole, i.e., not divided longitudinally into lobes (Figs 18A, C, E); median distal margin of first and second metatarsomeres without modified setae; cerci medially fused into a syncercus (Fig. 18G) Kinabalumyia Cerretti & Pape, gen. nov.
Mouthparts strongly reduced (proboscis very short, though palpus well developed). Male: proclinate orbital setae absent; postpedicel divided longitudinally into three lobes (Figs 4B; 6B); median distal margin of first and second metatarsomeres provided with a comb of flattened setae, which are distally jagged; cerci proximally fused, distally divided into two pointed branches (similar to the condition shown in Fig. 15F) Aporeomyia Pape & Shima
11 Vein M1 not reaching wing margin (i.e., ending freely in the wing membrane) (Fig. 9H–J) 12
Vein M1 reaching wing margin (Figs 9A, D, K, L; 10B, E, H–K) or fused to vein R4+5, so that cell r4+5 is petiolate (Figs 9E, F; 10C, F, G, L) 15
12 Male and female without proclinate orbital setae. Male: surstylus fused to epandrium Bezzimyia Townsend (in part)
Male and female with at least one proclinate orbital seta. Male: surstylus either freely articulating with or fused to epandrium 13
13 Prementum long and slender, at least as long as head height, and labella narrow. Postalar wall usually with at least one small setula. Postpronotum with two setae and facial profile not receding [Morinia lactineala (Pape), Moriniacarinata species group” (in part): Polleniidae]
Prementum shorter than head height, and labella fleshy and normally developed. Postalar wall bare. Facial profile receding or not; if not receding then postpronotum with three setae arranged in a right-angled triangle 14
14 Facial profile not receding, i.e., vibrissal angle distinctly in front of anterior margin of eye (Fig. 7E). Postpronotum with three setae arranged in a right-angled triangle (or nearly so) (as in Fig. 2C). Male: one proclinate orbital seta. Female: 2–3 proclinate orbital setae Oplisa Rondani (in part)
Facial profile receding, i.e., vibrissal angle approx. in line with or behind anterior margin of eye (head with postcranial surface oriented vertically) (Figs 7B, 8B–E). Postpronotum with 1–3 setae, if three then arranged in a line or an obtuse triangle (as in Fig. 2B). Male: at least two proclinate orbital setae. Female: 3–5 proclinate orbital setae 24
15 Posterior lappet of metathoracic spiracle distinctly larger than anterior lappet (Fig. 13H). Female: thorax yellow, brown or black 16
Anterior and posterior lappets of metathoracic spiracle approx. of equal size and standing out from spiracular rim (i.e., almost perpendicular to pleural surface) (Fig. 13I), sometimes lappets not differentiated at all (Fig. 13J). Female: thorax usually black or brown, rarely yellow 18
16 Vein R1 setose along full length dorsally. Vein R4+5 with setulae dorsally extending from base to approx. level of bend of vein M1. Cell r4+5 open (Fig. 14A, C). Parafacial bare. Facial ridge with a row of setae on lower 2/3 (Fig. 14A, D). Vibrissa well developed. Palpus absent. Female: head, thorax, legs and abdomen mostly pale yellow (Fig. 14). Male unknown Maurhinophora Cerretti & Pape, gen. nov.
Vein R1 bare. Vein R4+5 with a few short setulae confined at base or bare. Cell r4+5 open or petiolate. Parafacial setulose at least on upper half (Figs 6J; 13C, D). Facial ridge bare or with a few decumbent setae on lower third. Palpus well developed 17
17 Vein R4+5 with a few short setulae confined to base. Cell r4+5 petiolate (Fig. 9E, F). Anterior lappet of metathoracic spiracle without setae. Female: thorax yellow (Fig. 4G). Male: at least tip of wing membrane smoky (Fig. 9E); frons very narrow (i.e., head almost holoptic), without proclinate or lateroclinate orbital setae Baniassa Kugler
Vein R4+5 bare. Cell r4+5 open. Anterior lappet of metathoracic spiracle with setae. Female: thorax black or brown. Male: wing membrane evenly yellowish; frons broad with at least one proclinate or lateroclinate orbital seta (i.e., head non-holoptic) [Moriniacarinata species group” (in part): Polleniidae]
18 First postsutural supra-alar seta present and well developed, as long as or longer than notopleural setae (Fig. 13B, F) 19
First postsutural supra-alar seta absent or very short, distinctly shorter and weaker than notopleural setae (Fig. 13E, G) 24
19 Palpus absent. Facial ridge with a row of setae on lower 2/3 (Fig. 14D). Vein R1 entirely setulose dorsally. Vein R4+5 with setulae dorsally, extending from base to approx. level of bend of vein M1. Head, thorax, legs and abdomen mainly yellow (Fig. 14) Maurhinophora Cerretti & Pape, gen. nov.
Palpus present and usually well developed. Facial ridge with at most a few setulae just above vibrissa. Vein R1 bare. Vein R4+5 dorsally with a few short setulae confined to base or with one strong setula. Body colour variable 20
20 Palpus, tibiae, femora, pleural sclerites of thorax and sides of abdomen yellow (Fig. 5D). Head distinctly higher than long in lateral view. Facial ridge nearly straight. Lunule bare. Abdominal tergites 3 and 4 with strong, erect median discal setae (Fig. 11G). Male tergite 6 not differentiated Queximyia Crosskey
Palpus, tibiae, femora, pleural sclerites of thorax and sides of abdomen black or blackish-brown. Head varying from longer than high to higher than long. Facial ridge usually concave. Lunule bare or with setulae. Abdominal discal setae, if present, not as above. Male tergite 6 always present 211
21 Lunule with setulae (Fig. 13C). Katepimeron with at least one seta anteriorly. Male with or without orbital setae (lateroclinate and/or reclinate) Phyto Robineau-Desvoidy2
Lunule bare. Katepimeron bare. Male with orbital setae (lateroclinate and/or reclinate) 22
22 Upper half of facial plate carinate (Fig. 13A) (sometimes only slightly so). Base of vein R4+5 dorsally bare. Postalar wall usually with a tuft of short setulae (rarely bare). Anterior lappet of metathoracic spiracle usually with 1–7 setae. Prementum varying from normally developed to long and slender with narrow labella [Moriniacarinata species group” (in part): Polleniidae]
Facial plate not carinate. Base of vein R4+5 dorsally with one long setula (Fig. 10B). Postalar wall bare. Anterior lappet of metathoracic spiracle without setae. Prementum distinctly shorter than head height and labella unmodified (i.e., broad) 23
23 Facial profile moderately receding, i.e., vibrissal angle in line with antennal insertion (postcranial surface oriented vertically) (as in Fig. 7D). Male with orbital setae (only one, strong) Comoromyia Crosskey
Facial profile not receding, i.e., vibrissal angle produced forwards and in front of antennal insertion (Fig. 7I). Male without proclinate orbital setae Rhinomorinia Brauer & Bergenstamm3
24 Antenna distinctly shorter than height of gena (Figs 7B–D, F; 8A, B). Gena at least 0.5 times as high as compound eye; if less, then (i) mouthparts strongly reduced, vestigial and (ii) postangular section of vein M1 absent (Figs 4K; 9H–J). Male with proclinate orbital setae 25
Antenna at least as long as height of gena (usually distinctly longer), and gena at most 0.35 times as high as compound eye; if more, then mouthparts normally developed and vein M1 complete 29
25 Cell r4+5 open or vein M1 vanishing on wing membrane (i.e., postangular section of vein M1 absent) 26
Cell r4+5 long petiolate; petiole 0.8–2.5 times as long as postangular section of vein M1 (Figs 4L; 10L) 32
26 Postangular section of vein M1 absent and vein ending freely in wing membrane (Fig. 4K). Mouthparts strongly reduced, vestigial (Fig. 7B, C). Scutellum with only one pair of defined marginal setae (subapicals), diverging or subparallel. Thorax and abdomen characterised by a thick cover of silver microtomentum, which is brilliant in anterodorsal view (Figs 4K; 11A) (M. argyriventris, M. basilewskyi), or dull black (M. asetosa). Lunule bare. Male: trichia on arista bottle-brush like Melanophora Meigen4
Vein M1 complete, i.e., reaching wing margin (Figs 9L; 10J; 15E). Mouthparts small, but normally developed. Scutellum usually with two pairs of well-developed marginal setae (basals and converging or crossed apicals). Microtomentum of thorax and abdomen not as described above. Lunule bare or with setulae. Male: trichia on arista usually not bottle-brush like (except Marshallicona Cerretti & Pape, gen. nov.) 27
27 Male and female without proclinate orbital setae (Fig. 8A). Wing infuscate with 3 whitish, hyaline spots (Figs 5J; 10J) Trypetidomima Townsend
Male and female with at least one proclinate (or lateroclinate) orbital seta (Figs 7F; 15C). Wing membrane not as described above 28
28 Vein R1 dorsally setose on distal 1/4. Vein CuA+CuP not reaching wing margin. Costal sector cs5 longer than costal sector cs2 (Fig. 15E). Thorax blackish brown (Fig. 15A). Katepimeron bare. Lunule bare. Male: arista bottle-brush like; 4–5 proclinate orbital setae (Fig. 15C, D) Marshallicona Cerretti & Pape, gen. nov. (live habitus Fig. 1C)
Vein R1 dorsally bare. Vein CuA+CuP reaching wing margin. Costal sector cs5 shorter than costal sector cs2 (Fig. 9L). Thorax mainly yellow (Fig. 5B). Katepimeron usually with a few setulae anteriorly. Lunule usually setulose. Male: arista bare; 1–2 proclinate orbital setae Parazamimus Verbeke
29 Vein R1 entirely setulose dorsally. Scutellum with only one pair of defined marginal setae (subapicals), diverging or subparallel. Head shape strongly modified especially in male, with sunken face and vibrissal angle conspicuously projected forward and turned inwards apically (Fig. 7L). [Wing infuscate with 3 or 4 whitish spots (as in Fig. 10J). Male: proclinate orbital setae present; first aristomere elongated, more than twice as long as wide.] Shannoniella Townsend
Vein R1 bare dorsally, if with 1–3 setulae distally, then scutellum with at least 3 marginal setae. Head shape not as described above 30
30 Facial profile receding, i.e., vibrissal angle approx. in line with or behind anterior margin of eye (head with postcranial surface oriented vertically) (Figs 6A, K, L; 7B–D, F, G; 8A, C) 31
Facial profile not receding, i.e., vibrissal angle distinctly in front of anterior margin of eye (Figs 6G; 7A, E, I, J, K) 41
31 Cell r4+5 long petiolate (Figs 4L; 5C, G, L; 10C, F, G, L) 32
Cell r4+5 open (Figs 9A, D, G, K, L; 10B, E, J, K), closed at wing margin (Fig. 10H, I) or vein M1 ending freely in membrane (Fig. 9J) 34
32 Parafacial with a row of long robust setae (Fig. 8E). Male: arista not bottle-brush like. Female: wing without whitish posterior subapical spot Ventrops Crosskey (V. stuckenbergi Cerretti & Pape)
Parafacial with fine setulae at most on upper half (Fig. 13D). Male: arista bare, plumose or bottlebrush-like. Female: whitish posterior subapical spot on wing membrane absent or present 33
33 Both sexes with 3–7 proclinate orbital setae. Male: trichia on arista bottlebrush-like (Figs 7B; 13D); posterior margin of sternite 5 with a deep median cleft. Female: wing membrane mainly brownish with whitish posterior subapical spot Melanophora5
Male with 0–2, female with 1–3 proclinate orbital setae. Male arista bare to short pubescent, not bottlebrush-like; sternite 5 with almost straight posterior margin. Female: wing varying from hyaline to variously patterned but without whitish posterior subapical spot Paykullia Robineau-Desvoidy
34 Facial ridge with robust setae on lower 1/2 or more. Vein CuA+CuP reaching wing margin. Female: postpedicel with a row of setae medially or dorsally (Fig. 6L) Malayia Malloch
Facial ridge with only a few decumbent setulae reaching at most the lower 1/3. Vein CuA+CuP usually not reaching wing margin (except in Bixinia winkleri Cerretti, Lo Giudice & Pape from Australia). Female: postpedicel without setae 35
35 Parafacial bare. Lateral vertical seta not differentiated from postocular row (as in Fig. 8A) 36
Parafacial setose at least on upper half (Figs 6A; 8E). Lateral vertical seta well differentiated from postocular row (Fig. 8D, E); if not differentiated, then lunule with setulae 38
36 Bend of vein M1 distinctly rounded and very close to wing margin (Fig. 9G). Antenna distinctly longer than compound eye height, and facial ridge longer than frons (Fig. 6K) Bixinia Cerretti, Lo Giudice & Pape
Bend of vein M1 well removed from wing margin (Fig. 9K, L). Antenna shorter than compound eye height and facial ridge shorter than frons (Fig. 8A, C) 37
37 Male and female with at least 2 proclinate orbital setae (Fig. 8C). Wing infuscate mostly along veins, without whitish spots. Lower calypter without long, blackish setulae along margin. Posterior margin of eye indented in lateral view (Fig. 8C). Eye enormously developed so that gena and parafacial are practically obliterated. Male cerci very short Ventrops Crosskey6
Male and female without proclinate orbital setae (Fig. 8A). Wing infuscate with 3 whitish, hyaline spots (Figs 10J). Lower calypter with long, blackish setulae along margin. Posterior eye margin not indented (Fig. 8A). Eye large but not enormously developed so that both gena and parafacial are distinct. Male cerci well developed Trypetidomima Townsend
38 Antennal insertion distinctly above eye middle. Antenna long, at least as long as eye height. Second aristomere elongated, 2–3 times as long as wide Acompomintho Villeneuve
Antennal insertion at or below eye middle (Figs 6A; 8D, E). Antenna distinctly shorter that eye height. Second aristomere at most as long as wide 39
39 Male and female without proclinate (or lateroclinate) orbital setae (Fig. 6A). Lunule setose. Legs largely red (Fig. 4A). Male: surstylus fused to epandrium Apomorphyto Cerretti, Lo Giudice & Pape
Male and female with proclinate orbital setae (Fig. 8D, E). Lunule bare. Legs black or blackish-brown. Male: surstylus not fused to epandrium 40
40 Abdominal tergites extensively covered with whitish-grey reflecting microtomentum except along posterior margins of tergites 3–5 and along a narrow longitudinal median vitta. Head and thorax extensively covered with grey reflecting microtomentum. Basicosta yellow. Male: arista pubescent, longest trichia distinctly longer than maximum, basal diameter or arista. Body length: 6 mm Ventrops Crosskey (undescribed species from South Africa)
Head, thorax and abdomen shiny black or nearly so, i.e., with microtomentum sparse or almost absent. Basicosta black or yellow. Male: arista apparently bare, i.e., trichia clearly shorter than its maximum diameter. Body length less than 4 mm Ventrops Crosskey7
41 Distal section of CuA+CuP (i.e., distal to junction with CuA) approx. 1/2 of total length of CuA+CuP (Fig. 9C, D; 10A). Anterior katepisternal seta weak and at most 2/3 as long as posterior one 42
Distal section of CuA+CuP approx. 3/4 of total length of CuA+CuP (Figs 9A, E, G; 10E–G). Anterior katepisternal seta robust and more than 2/3 as long as posterodorsal katepisternal seta 43
42 Anterior katepisternal seta more than 1/2 as long as posterior seta. Second aristomere slightly elongated, approx. 1.5 times as long as wide (Fig. 6G, H). Male: fore tarsus not elongated or compressed (Fig. 4F); wing membrane hyaline or evenly smoked (Fig. 9D) Azaisia Villeneuve
Anterior katepisternal seta distinctly less than 1/2 as long as posterior seta. Second aristomere at most as long as wide (Fig. 7A). Male: fore tarsus greatly elongated and laterally compressed (Fig. 4J); wing membrane darkened around distal third of vein R2+3 (Fig. 4J) Macrotarsina Schiner
43 Setae present on lunule (Fig. 13C), notopleuron and katepimeron simultaneously. Median process of ventral sclerotisation of distiphallus interrupted proximally and not connected to ventral plate (Fig. 12G) Phyto Robineau-Desvoidy (P. pauciseta Herting)
Setae never present on lunule, notopleuron and katepimeron simultaneously. Median process of ventral sclerotisation of distiphallus not interrupted, running from ventral plate to tip of phallus 44
44 Postpronotum with three setae arranged in right-angled triangle or nearly so; if with two setae (one species of Stevenia), then parafacial with strong setae in lower half 45
Postpronotum with three setae arranged in a line or in a shallow triangle; if with only two setae, then parafacial bare 47
45 Parafacial bare or with hair-like setulae in upper half; if parafacial setae continue on lower half, these are never robust (Fig. 7E). Cell r4+5 never petiolate Oplisa Rondani (in part)
Parafacial with a row of strong bristly setae in lower half (Fig. 7J, K). Cell r4+5 narrowly open, closed at wing margin or petiolate (Fig. 10H, I, L) 46
46 Cell r4+5 narrowly open, closed at wing margin or, more rarely, short-petiolate (petiole, when present, shorter than crossvein r-m). Mid tibia with 1 anterodorsal seta. Male habitus in lateral view as in Fig. 5H Tricogen a Rondani
Cell r4+5 distinctly petiolate (Fig. 10L) (petiole approx. as long as crossvein r-m in S. acutangula). Mid tibia with at least 2 anterodorsal setae. Male habitus in lateral view as in Fig. 5G Stevenia Robineau-Desvoidy
47 Arista plumose, i.e., longest trichia approx. 2.5 times as long as maximum diameter of arista (Fig. 8B) Tromodesia Rondani
Arista bare or with trichia at most 1.5 times as long as maximum diameter of arista 48
48 Base of vein R4+5 bare or with a few, short, fine setulae, distinctly shorter than crossvein r-m. Vibrissal angle more or less in line with antennal insertion (postcranial surface oriented vertically) (Figs 6G; 7E) 49
Base of vein R4+5 with at least one strong setula, longer than crossvein r-m, with or without additional shorter setulae. Vibrissal angle produced forwards and in front of antennal insertion (Figs 7I; 12C) 51
49 Cell r4+5 open or closed at wing margin, very rarely petiolate but petiole at most approx. half as long as crossvein r-m 50
Cell r4+5 with a petiole 1.2–2.5 times as long as crossvein r-m 52
50 Frontal vitta, palpus (Figs 6G, H; 7A) and basicosta yellow 42
Frontal vitta, palpus and basicosta black Metoplisa Kugler
51 Cell r4+5 open or closed at wing margin, rarely petiolate with petiole approx. half as long as crossvein r-m. Scutum with two presutural microtomentose vittae. Palpus at least as long as postpedicel (Fig. 7I). Male: head not holoptic, with upper reclinate orbital setae usually differentiated (except R. capensis)] Rhinomorinia Brauer & Bergenstamm (in part)
Cell r4+5 with a petiole 1.2–2.5 times as long as crossvein r-m. Scutum dark with no presutural microtomentose vittae differentiated. Palpus varying from shorter to longer than postpedicel. Male: head holoptic or not so, with or without upper reclinate orbital setae 52
52 Parafacial entirely bare. Palpus shorter than postpedicel. Lunule bare. Male: head almost holoptic, upper reclinate orbital setae not differentiated (Fig. 12A, C). Male terminalia as in Fig. 12D, E Melanomyoides Crosskey
Parafacial with a row or short setulae. Palpus longer than postpedicel. Lunule with setulae. Male: head not holoptic, with upper reclinate orbital setae usually differentiated Rhinophora Robineau-Desvoidy
Figure 4. 

Rhinophoridae, habitus in lateral view. A Apomorphyto inbio ♂ (Costa Rica) [holotype] B Aporeomyia sp. ♂ (Malaysia, Sabah) C Axinia zantae ♂ (Australia) D Axinia arenaria ♂ (Australia) E Axinia brevispica ♂ (Australia) F Azaisia sp. ♂ (Portugal, Azores) G Baniassa fascipennis ♀ (Israel) [paratype] H Bezzimyia hansoni ♀ (Costa Rica) I Comoromyia sp. ♀ (Madagascar) J Macrotarsina longimana ♂ (Italy) K Melanophora basilewskyi ♂ (Kenya) L Melanophora roralis ♂ (Italy).

Figure 5. 

Rhinophoridae, habitus in lateral view. A Oplisa tergestina ♂ (Italy) B Parazimimus congolensis ♂ (Burundi) C Paykullia partenopea ♂ (Italy) D Queximyia flavipes ♂ (South Africa) E Rhinodonia antiqua ♂ (New Caledonia) [holotype] F Rhinomorinia sarcophagina ♂ (Italy) G Stevenia palermitana ♂ (Italy) [holotype] H Tricogena rubricosa ♂ (Morocco) I Tromodesia angustifrons ♀ (Israel) [paratype] J Trypetidomima lutea ♀ (Brazil) K Ventrops incisus ♂ (Tanzania) [paratype] L Ventrops stuckenbergi ♂ (Namibia) [holotype].

Figure 6. 

Rhinophoridae, head in lateral view. A Apomorphyto inbio ♂ (Costa Rica) [holotype] B Aporeomyia sp. (Malaysia, Sabah) C Axinia arenaria ♂ (Australia) D Axinia brevispica ♂ (Australia) E Axinia lucaris ♂ (Australia) F Axinia zentae ♂ (Australia) G, H Azaisia sp. ♂ (Portugal, Madeira) G lateral view, H dorsolateral view I Bezzimyia bisecta ♂ (Costa Rica) J Baniassa fascipennis ♀ (Israel) [paratype] K Bixinia collessi ♂ (Australia) [paratype] L Malayia fuscinervis ♀ (Malaysia, Malay Peninsula).

Figure 7. 

Rhinophoridae, head in lateral view. A Macrotarsina longimana ♂ (Italy) B, C Melanophora basilewskyi ♂ (Kenya) B lateral view C fronto-lateral view D Melanophora roralis ♂ (Italy) E Oplisa tergestina ♂ (Italy) F Parazamimus congolensis ♂ (Burundi) G Paykullia cf. nubilipennis ♂ (Italy) H Rhinodonia antiqua ♂ (New Caledonia) [holotype] I Rhinomorinia sarcophagina ♂ (Italy) J Stevenia palermitana ♂ (Italy) [paratype] K Tricogena rubricosa ♂ (Morocco) L Shannoniella setinervis ♂ (Brazil).

Figure 8. 

Rhinophoridae, head in lateral view. A Tripetidomima lutea ♂ (Brazil) B Tromodesia angustifrons ♀ (Israel) [paratype] C Ventrops milichioides ♂ (Tanzania) D Ventrops hannemariae ♀ (Tanzania) E Ventrops stuckenbergi ♂ (Namibia) [holotype].

Figure 9. 

Rhinophoridae, wing. A Apomorphyto inbio ♂ (Costa Rica) [holotype] B Aporeomyia sp. (Malaysia, Sabah) ♂ C Axinia cf. brevispica ♀ (Australia) D Azaisia sp. ♂ (Portugal, Madeira) E, F Baniassa fascipennis (Israel) E ♂ [paratype] F ♀ [paratype] G Bixinia collessi ♂ (Australia) [paratype] H Bezzimyia bisecta ♂ (Costa Rica) I Bezzimyia busckii ♂ (Costa Rica) J Melanophora basilewskyi ♀ (Kenya) K Metoplisa carbonaria ♂ (Israel) L Parazamimus congolensis ♂ (Burundi).

Figure 10. 

Rhinophoridae, wing. A Oplisa aterrima ♂ (Italy) B Oplisa tergestina ♂ (Italy) C Paykullia partenopea ♂ (Italy) D Rhinodonia antiqua ♂ (New Caledonia) [holotype] E Rhinomorinia sarcophagina ♂ (Italy) F Rhinophora lepida ♂ (Italy) G Stevenia etrusca ♀ (Italy) [paratype] H Tricogena rubricosa ♀ (Morocco) I Tromodesia angustifrons ♀ (Israel) [paratype] J Trypetidomima lutea ♂ (Brazil) K Ventrops milichioides ♂ (Tanzania) L Ventrops stuckenbergi ♂ (Namibia) [holotype] M Rhinopeza gracilis ♂ (Papua New Guinea) [holotype] N Polleniidae, wing: Morinia carinata ♂ (South Africa).

Figure 11. 

Rhinophoridae, abdomen. A Melanophora basilewskyi ♂ (Kenya) B Oplisa tergestina ♂ (Italy) C Parazamimus congolensis ♂ (Burundi) D Rhinomorinia sarcophagina ♂ (Italy) E Tromodesia angustifrons ♀ (Israel) [paratype] F Ventrops stuckenbergi ♂ (Namibia) [holotype] G Queximyia flavipes ♂ (South Africa). A–F dorsal view G lateral view.

Figure 12. 

Melanomyoides capensis (South Africa). A Habitus in lateral view B abdomen in dorsal view C head in lateral view D, E epandrial complex in posterior view (D) and lateral view (E) F, G Phyto adolescens (Italy), hypandrial complex in posterior view (F) and lateral view (G).

Figure 13. 

SEM images of diagnostic characters. A, B Polleniidae A Morinia carinata ♂ (South Africa) [paratype], detail of head in anterodorsal view B Morinia carinata ♂ (South Africa) [paratype], thorax in dorsolateral view [colour coding as E–G] C–N Rhinophoridae C Phyto adolescens ♂ (Italy), detail of head in anterodorsal view D Melanophora chia ♂ (Italy), detail of anterior part of head in lateral view E–G thorax in dorsolateral view [B, E–G colour coding: red = first postsutural supra-alar seta; yellow = second postsutural supra-alar seta; blue = third postsutural supra-alar seta; green = posterior notopleural seta] E Parazamimus congolensis ♀ (Burundi) F Queximyia flavipes ♂ (South Africa) G Rhinomorinia sarcophagina ♂ (Italy) H–J metathoracic spiracle [colour coding: green = anterior lappet; red = posterior lappet] H Baniassa fascipennis ♀ (Israel) I Rhinomorinia sp. ♀ (South Africa) J Melanophora basilewskyi ♀ (Kenya) K–N Axinia spp. (Australia), female oviscapt K, L Axinia zentae in lateral view (K) and posterior view (L) M, N Axinia sp. in lateral view (M) and posterior view (N).

New taxa

Four new genera are here erected to accommodate five new species, which do not fit within any of the current generic concepts within Rhinophoridae according to our phylogenetic analysis (see Fig. 20, and “Phylogeny and suprageneric classification” section below). Erecting these new genera is considered superior to alternatives involving generic lumping, as it will facilitate our communication and visualisation of the ever-increasing morphological diversity of the world rhinophorids.

Afrotropical region

Maurhinophora Cerretti & Pape, gen. nov.

Fig. 14

Unambiguous character state changes

(Table 1, Fig. 20). Global apomorphies: none; local apomorphies: 13:1, 14:1, 21:2; 30:2, 42:1, 44:1.

Diagnosis

Head : head higher than long in lateral view. Facial ridge 1.1 times as long as frons. Ocellar setae virtually absent. Frons approx. 0.9 times as wide as compound eye in dorsal view. Median (= inner) vertical setae strong and crossed. Five medially crossed frontal setae, slightly reclinate, descending to approx. half level of pedicel. Fronto-orbital plate with some short setulae. Two proclinate orbital setae. One upper lateroclinate orbital seta. Parafacial bare, at its narrowest point at most 1.5 times as wide as maximum diameter of arista. Vibrissal angle receding. Vibrissa well developed, arising slightly below level of lower facial margin. Lower facial margin sunken and not visible in lateral view. Facial ridge slightly and evenly convex with a row of setae on lower 2/3, decreasing in size dorsally. Face deeply concave, antennae hidden in lateral view. Antenna long and narrow, much longer than height of gena. Postpedicel narrowly elongated approx. 5 times as long as pedicel. Arista bare (or apparently so). Arista thickened in proximal 2/5–1/2; second aristomere at most as long as wide. Lunule bare. Gena, in profile, approx. 1/5 as high as compound eye. Palpus absent.

Thorax : prosternum bare. Postpronotum with three setae arranged in triangle. Three postsutural supra-alar setae (first postsutural supra-alar seta well developed, i.e., longer than posterior notopleural seta and approx. the same size as anterior notopleural seta). Scutellum with one pair of well-developed basal setae and one pair of strong, horizontal and crossed apical setae; basal setae placed at level of apical setae. Anatergite with a tuft of short setulae below lower calypter. Subscutellum moderately swollen, not fully sclerotised. Posterior lappet of metathoracic spiracle larger than anterior lappet (as in Baniassa). Lower calypter distinctly tongue-shaped (ground-plan trait of Rhinophoridae) (Fig. 2E). Costal sector cs2 setose ventrally. Costal spine not differentiated from general costal setae. Costal sector cs5 clearly shorter than costal sector cs2. Vein R1 entirely setulose dorsally. Vein R4+5 with setulae dorsally extending from base to approx. level of bend of vein M1. Bend of vein M1 well developed, rounded and well removed from wing margin. Crossvein dm-m forming a right angle with proximal section of M4. Vein CuA+CuP not reaching wing margin. Preapical anterodorsal seta of fore tibia longer than preapical dorsal seta. Fore tarsus not compressed. Tibiae of mid and hind leg normally developed. Mid tibia with one, short subdistal anterodorsal seta. Hind tibia with 3 dorsal preapical setae.

Abdomen : tergites without microtomentum and with relatively strong and suberect general setulae; syntergite 1+2 without median discal setae, tergite 3 with one pair of strong median discal setae, tergites 4 and 5 with a row of strong marginal setae (discal setae not differentiated).

Distribution

Afrotropical – Mauritius.

Type species

Maurhinophora indoceanica Cerretti & Pape, sp. nov., by present designation.

Etymology

The generic name is a composite word formed from the first part of the name of the island Mauritius, to which the known species is restricted, and the name Rhinophora, which is the type-genus for the family-group name Rhinophoridae. The name should be treated as a feminine noun.

Figure 14. 

Maurhinophora indoceanica gen. et sp. nov. ♀ (Mauritius) [holotype] A habitus in lateral view B abdomen in dorsal view C wing in dorsal view D head in lateral view.

Maurhinophora indoceanica Cerretti & Pape, sp. nov.

Fig. 14

Type material

Holotype ♀: Mauritius; /Corps de la Garde, to 2,200’ /4.vi.1971 /A.M. Hudson /B.M. 1971-346. (NHMUK).

Description

Female. Body length: 5.5 mm. Colouration: head, thorax (including tegula, basicosta, wing veins and legs) yellow; abdomen mostly yellow with blackish brown transversal bands on posterior 1/3 of syntergite 1+2, and posterior 1/2 of tergites 3–5. Head: frontal vitta as wide as fronto-orbital plate. Two strong proclinate orbital setae (one additional short proclinate orbital seta is present posterior to the main ones). Parafacial narrow with only one setula below lower frontal seta. Lateral (= outer) vertical seta well developed though not strongly differentiated from strongest uppermost postocular setae. Prementum approx. 2 times as long as wide; labella not elongated, normally developed. Thorax: one posthumeral seta (medial); 1 + 3 supra-alar setae (posterior postsutural supra-alar weak); 0 + 2 intra-alar setae; 2(3) + 3 dorsocentral setae; 0 + 1 acrostichal setae. Two strong, diverging katepisternal setae. Legs: fore tibia with 1 posterior seta. Abdomen: mid-dorsal depression on syntergite 1+2 confined to anterior half.

Distribution

Afrotropical – Mauritius.

Etymology

The species epithet is derived from the name of the Indian Ocean and should be treated as a Latin adjective.

Remarks

We consider the description of a new species based on a single female as warranted due to the quite remarkable habitus and the occurrence on a small oceanic island, which means that the likelihood of complications due to the lack of male material can be considered low.

Neotropical region

Marshallicona Cerretti & Pape, gen. nov.

Figs 1C, 15

Unambiguous character state changes

(Table 1, Fig. 20). Global apomorphies: none; local apomorphies: 2:1, 9:1, 43:1, 51:1, 52:1, 56:1, 66:0, 68:1.

Diagnosis

Head : head higher than long in lateral view. Facial ridge 0.4 times as long as frons. Ocellar setae virtually absent. Frons approx. 0.8 times as wide as compound eye in dorsal view. Median vertical setae strong, subparallel. Six to eight medially crossed frontal setae, slightly reclinate, descending to level of upper margin of scape. Fronto-orbital plate with few short setulae confined to upper half. Four or five proclinate orbital setae (posterior two slightly lateroclinate). One weak upper lateroclinate orbital seta (usually not distinguishable from uppermost frontal setae). Parafacial bare, at its narrowest point approx. as wide as width of postpedicel. Vibrissal angle receding. Vibrissa well developed, arising at level of lower facial margin. Lower facial margin not sunken though not visible in lateral view. Facial ridge concave with decumbent setulae on lower 1/3–2/5. Face slightly concave, antennae not hidden from view in profile. Antenna approx. as long as height of gena. Postpedicel sub-ovoid, approx. 1.5 times as long as pedicel. Arista bottlebrush-like, trichia longer that maximum diameter of arista. Arista thickened in proximal 1/5 or less; second aristomere at most as long as wide. Lunule hidden by inner anterior margins of fronto-orbital plate. Gena, in profile, approx. 1/2 as high as compound eye. Palpus stout, clavate, with a few thin setulae on apical 1/4.

Thorax : prosternum bare. Postpronotum with two setae. One postsutural supra-alar seta (i.e., first and third post sutural supra-alar setae absent). Scutellum with one pair of well-developed basal setae and one pair of strong, horizontal and crossed apical setae; basal setae placed dorsally with respect to apical setae. Anatergite bare. Subscutellum moderately swollen, not fully sclerotised. Metathoracic spiracular lappets virtually absent. Lower calypter distinctly tongue-shaped (ground-plan trait of Rhinophoridae) (Fig. 2E). Costal sector cs2 setose ventrally. Costal spine not differentiated from general costal setae. Costal sector cs5 longer than costal sector cs2 (Fig. 15E). Vein R1 dorsally setose on distal 1/4. Base of R4+5 entirely bare. Bend of vein M1 shallow, well removed from wing margin. Crossvein dm-m forming an acute angle with proximal section of M4. Vein CuA+CuP not reaching wing margin. Preapical anterodorsal seta of fore tibia approx. as long as preapical dorsal seta. Fore tarsus not compressed. Tibiae of mid and hind leg normally developed. Mid tibia with one anterodorsal seta. Hind tibia with 3 dorsal preapical setae.

Abdomen : slightly elongated, virtually without microtomentum and without distinct marginal and discal setae.

Male terminalia (Fig. 15F–H): posterior margin of sternite 5 with a deep median notch; lateral lobe rounded posteriorly. Tergite 6 bare, medially divided into two hemitergites; tergite 6 divided from syntergosternite 7+8 by a wide membrane. Connection between sternite 6 and syntergosternite 7+8 on right side membranous. Cerci well developed, not fused medially. Basal 1/3 of cerci convex and covered with short setae; distal 2/3 straight and narrowly digitiform (branches symmetrically diverging and well separated). Surstylus well developed, lobe-like in lateral view; lateral side of surstylus broadly convex at approx. mid length. Surstylus not fused to epandrium. Bacilliform sclerite firmly fused to laterobasal margin of surstylus. Hypandrial arms not fused medially. Connection between phallic guide and pregonite sclerotised. Postgonite without anterior seta. Epiphallus well developed and attached dorsomedially to basiphallus. Extension of dorsal sclerite of distiphallus entirely fused medially into a single sclerite and proximally fused to dorsal sclerite of distiphallus. Median process of ventral sclerotisation of distiphallus present, divided medially into two hemisclerites, which are both proximally fused to ventral plate of distiphallus. Acrophallus simple and scale-like spinules present lateroventrally.

Distribution

Neotropical – Ecuador.

Type species

Marshallicona quitu Cerretti & Pape, sp. nov., by present designation.

Etymology

The generic name is a composite word formed from the name of our colleague and friend Steve Marshall, who collected the type series and took the photo of a living specimen (Fig. 1C), and from the Latin noun ‘icona’, meaning image, in honour of Steve’s remarkable skills in natural history photography. The name should be treated as a feminine noun.

Figure 15. 

Marshallicona quitu gen. et sp. nov. (Ecuador) ♂. A Habitus in lateral view B abdomen in dorsal view C head in lateral view D head in frontal view E wing in dorsal view F, G epandrial complex in posterior view (F) and lateral view (G) H hypandrial complex in lateral view [A–D holotype, E–H paratype].

Marshallicona quitu Cerretti & Pape, sp. nov.

Figs 1C, 15

Type material

Holotype ♂: Mindo-Bellavista Ecuador /Bellavista Cloud Forest Reserve /0°3'28.57"S, 78°46'6.02"W /2000m 1 May 2001 /S.A. Marshall and S.P.L. Luk (PUCE). Paratypes 2 ♂♂: same data as holotype (NHMD, MZUR).

Description

Male. Body length: 4.5–5.5 mm. Colouration: head mostly black in ground colour, antenna and palpus brown; occiput and posterior 2/3 of fronto-orbital plate shiny without microtomentum, anterior 1/3 of fronto-orbital plate and parafacial covered with silver reflecting microtomentum; thorax (including tegula, basicosta, wing veins and legs) black in ground colour, microtomentum virtually absent; abdomen mainly yellow except brownish posterior margin of tergites 1+2, 3 and 4 and on posterior 1/2–2/3 of tergite 5. Wing membrane infuscate around veins. Head: frontal vitta slightly narrower than fronto-orbital plate (measured at midlength). Parafacial entirely bare below lower frontal seta. Lateral vertical seta not differentiated from strongest uppermost postocular setae. Prementum stout, not longer than wide; labella broad. Thorax: one posthumeral seta (medial); 1 + 1 supra-alar setae; 0 + 2 intra-alar setae; 2(3) + 3 dorsocentral setae; 0–3 + 1 acrostichal setae. One or 2 katepisternal setae. Legs: fore tibia without posterior seta. Abdomen: mid-dorsal depression on syntergite 1+2 confined to anterior half.

Distribution

Neotropical – Ecuador.

Etymology

The species epithet is derived from the Quitu tribe, the pre-Columbian indigenous people who founded the city of Quito, which is now the capital of Ecuador. The name should be treated as a noun in apposition.

Neotarsina Cerretti & Pape, gen. nov.

Figs 16, 17

Unambiguous character state changes

(Table 1, Fig. 20). Global apomorphies: 33:1; local apomorphies: 21:1, 59:1, 72:1.

Diagnosis

Head : head higher than long in lateral view. Facial ridge 0.6 times as long as frons. Ocellar setae virtually absent. Frons 0.3–0.5 (male), 0.8–0.9 (female) times as wide as compound eye in dorsal view. Median vertical setae converging or crossed, though very short, at most as long as antenna. Ten to 20 short, medioclinate frontal setae, descending to level of upper margin of scape. Fronto-orbital plate bare or with scattered setulae interspersed between frontal setae. Proclinate orbital setae absent. Upper reclinate orbital seta absent. Parafacial bare, at its narrowest point 0.8–1.2 times as wide as width of postpedicel. Vibrissal angle receding. Vibrissa weak, i.e., barely distinguishable from setae of subvibrissal ridge, arising at level of lower facial margin. Lower facial margin not sunken though not visible in lateral view. Facial ridge concave with decumbent setulae on lower 1/5–2/5. Face slightly concave, antennae not hidden in lateral view. Antenna shorter than height of gena. Postpedicel sub-ovoid, approx. 1.0–1.7 times as long as pedicel. Arista bare. Arista thickened on proximal 1/10–1/5 of its length; second aristomere at most as long as wide. Lunule bare. Gena, in profile, 2/5–1/2 as high as compound eye. Palpus very short 1–2 times as long as wide, bare.

Thorax : prosternum bare. Postpronotum with 2–3 setae. One postsutural supra-alar seta (i.e., first and third post sutural supra-alar setae absent). Scutellum with one pair of basal setae and one pair of, crossed, horizontal apical setae; basal setae placed dorsally with respect to apical setae. Anatergite bare. Subscutellum moderately swollen or flat, not fully sclerotised. Metathoracic spiracular lappets small, subequal in size and directed outwards. Lower calypter distinctly tongue-shaped (ground-plan trait of Rhinophoridae) (Fig. 2E). Costal sector cs2 usually setose ventrally. Costal spine not differentiated from general costal setae. Costal sector cs5 approx. as long as costal sector cs2. Vein R1 dorsally bare. Base of R4+5 entirely bare. Bend of vein M1 indistinct; i.e., M1 evenly curved forward without forming a distinct bend. Crossvein dm-m forming a right angle with proximal section of M4. Vein CuA+CuP not reaching wing margin. Preapical anterodorsal seta of fore tibia longer than preapical dorsal seta. Fore tarsus strongly laterally compressed in both sexes. Tibiae of mid and hind legs laterally compressed and distinctly keeled dorsally. Mid tibia without anterodorsal setae. Hind tibia with three dorsal preapical setae.

Abdomen : lightly elongated, varying from slightly microtomentose to virtually without microtomentum. Marginal and discal setae not differentiated from general setulae.

Male terminalia : posterior margin of sternite 5 with a deep median notch; lateral lobe rounded posteriorly. Tergite 6 plate-like, with median marginal setae; tergite 6 divided from syntergosternite 7+8 by a membrane. Connection between sternite 6 and syntergosternite 7+8 fused on right side. Cerci well developed, basally broad, narrowing toward apex and apically pointed, well separated medially. Surstylus well developed, sub-triangular in lateral view; lateral side of surstylus not or only slightly convex. Surstylus fused to epandrium. Bacilliform sclerite articulated (i.e., not fused) to laterobasal margin of surstylus. Hypandrial arms not fused medially. Connection between phallic guide and pregonite membranous. Postgonite without anterior seta. Epiphallus well developed and attached dorsomedially or dorsomedially to basiphallus. Extension of dorsal sclerite of distiphallus divided medially into two hemisclerites which are proximally not fused to dorsal sclerite of distiphallus. Median process of ventral sclerotisation of distiphallus present, not interrupted, i.e., running from the ventral plate to tip of phallus, and not divided medially. Acrophallus simple and scale-like spinules not differentiated.

Distribution

Neotropical – Peru, Trinidad and Tobago (Trinidad).

Type species

Neotarsina caraibica Cerretti & Pape, sp. nov., by present designation.

Etymology

The generic name is a composite word formed from the Latin word ‘neo’ meaning new, and ‘tarsina’ [from Latin ‘tarsus’ and Greek: ‘tarsos’, the flat part of a human foot] as in the last part of the genus-group name Macrotarsina, in reference to the modified, laterally-compressed fore tarsus characterising the two new species described below. The name should be treated as a feminine noun.

Figure 16. 

Neotarsina andina gen. et sp. nov. (Peru). A Female habitus in lateral view B male abdomen in dorsal view C female abdomen in dorsal view D, E male head in lateral view (D) and frontal view (E) F, G female head in lateral view (F) and frontal view (G) H female fore tibia and tarsus I male wing in dorsal view J–L epandrial complex in posterior view (J) and lateral view (K) L hypandrial complex in lateral view [B, D–E, I–L holotype; A, C, F, G paratype].

Figure 17. 

Neotarsina caraibica gen. et sp. nov. (Trinidad and Tobago). A Male habitus in lateral view B female habitus in lateral view C male abdomen in dorsal view D female abdomen in dorsal view E female fore tibia and tarsus F, G male head in lateral view (F) and frontal view (G) H, I female head in lateral view (H) and frontal view (I) view J male wing in dorsal view K–M mpandrial complex in posterior (K) and lateral (L) view M hypandrial complex in lateral view [A, C, F–G, J holotype; B, D–E, H–I, K–M paratypes].

Neotarsina andina Cerretti & Pape, sp. nov.

Fig. 16

Type material

Holotype ♂: Peru: Raymondi /monoculture (loc. 11) /6°45'21"S, 79°51'05"W /J. Krausová, 15.v.09 (CULSP). Paratype ♀: PERU: Pimental, 165 m /agroforest (loc. 81) /8°31'30"S, 74°46'30"W /J. Krausová, 1.xi.10 (CULSP).

Notes on type specimens

The holotype lacks postpronotal and scutellar setae but is otherwise in good condition. The paratype is in fair general condition but lacks most of the setae on the thorax.

Description

Male. Body length: ca. 4 mm. Colouration: head mostly black except antenna and palpus which are pale yellow, parafacial, gena and lower occiput covered with dense silver reflecting microtomentum; fronto-orbital plate polished shiny black, except narrowing microtomentose along medial margin; thorax entirely black or dark brown in ground colour; thoracic pleura covered with silver reflecting microtomentum; femora mostly yellow except dark brown proximally, tibiae and tarsi dark brown; calypters whitish; tegula and basicosta pale yellow; wing membrane hyaline; halter yellow; abdomen black in ground colour, tergites 3 and 4 laterally covered with silver reflecting microtomentum. Head: frontal vitta measured at mid-length approx. 1.7 times as wide as fronto-orbital plate at same level. Postpedicel 1.4–1.7 times as long as pedicel. First and second aristomeres approx. as long as wide. Frontal setae medioclinate, slightly proclinate near antennal insertion. Median vertical setae short, approx. 0.2 times as long as compound eye height. Parafacial approx. 1.2 times as wide as postpedicel. Facial ridge bare or with few short setae above vibrissa. Gena, in profile, 2/5 as high as compound eye. Frons 0.5 times as wide as eye in dorsal view. Prementum stubby, approx. 2 times as long as wide; labella not elongated, normally developed. Palpus short, clavate. Thorax: two postpronotal setae; two posthumeral setae; 0 + 1 supra-alar setae (i.e., first and third postsutural supra-alar setae absent); 0 + 2 intra-alar setae; 2 + 3 dorsocentral setae (first presutural dorsocentral barely distinguishable from general setulae); one presutural acrostichal seta; postsutural acrostichal setae apparently not differentiated. Fore tibia approx. 1.5 times as long as first tarsomere. Costal sector cs2 nearly bare ventrally. Abdomen: Tergite 5 very short, approx. 0.5–0.6 times as long as tergite 4. Male terminalia: Surstylus narrowly triangular in lateral view. Cerci relatively narrow with a clear, though shallow, bottleneck restriction at approx. mid length; apical 1/5 of cerci strongly narrowing and pointed. Female. Body length: 5 mm. Female differs from male as follows: Head: fronto-orbital plate entirely shiny and parafacial with less dense microtomentum. Frontal vitta measured at mid-length approx. 2.0 times as wide as fronto-orbital plate at same level. Frons 0.8 times as wide as eye in dorsal view.

Distribution

Neotropical – Peru.

Etymology

The species epithet is derived from the name of the Andes mountain range and should be treated as a Latin adjective.

Neotarsina caraibica Cerretti & Pape, sp. nov.

Fig. 17

Type material

Holotype ♂. TRINIDAD: Curepe /Santa Margarita /malaise trap /12–14.vi.1972 //B.R. Pitkin coll. /BMNH(E) 1997-41 (NHMUK). Paratypes: 1 ♂: same data and repository as holotype; 1 ♀: Simla, Trin. [Trinidad and Tobago] /II-16-1966 /W. D. Duckworth (USNM, unique specimen identifier: USNMENT01519745).

Description

Male. Body length: ca. 4.5–5.5 mm. Colouration: head mostly dark brown, except antenna and palpus which are yellow and genal dilation, which is brownish; head evenly covered with grey reflecting microtomentum. Scutum brown with three or four pale longitudinal vittae; thoracic pleura covered with silver reflecting microtomentum; tegula, basicosta light brown, calypters and wing membrane slightly infuscate; halter yellow; coxae and femora yellow; tibiae brown; tarsi dark brown; abdomen yellowish to light brown, weakly microtomentose. Head: frontal vitta 0.5–0.7 times as wide as fronto-orbital plate, both measured at approx. midlength. Ocellar triangle setulose, without ocellar setae. Postpedicel approx. 1.0–1.2 times as long as pedicel. Arista thickened in proximal 1/8–1/10; first and second aristomere approx. as long as wide. Frontal setae medioclinate, slightly proclinate near base of antenna. Median vertical setae short, 0.2–0.3 times as long as height of compound eye, crossed medially. Parafacial 0.8–0.9 times as wide as postpedicel. Facial ridge with few short setae above vibrissa. Gena, in profile, 2/5–1/2 as high as compound eye. Frons approx. 0.3 times as wide as compound eye in dorsal view. Prementum stubby, 1–2 times as long as wide; labella not elongated, normally developed; palpus exceptionally reduced. Thorax: two postpronotal setae; one posthumeral seta; 0 + 1 supra-alar setae (i.e., first and third postsutural supra-alar setae absent); 1 + 1–2 intra-alar setae; 2 + 3 dorsocentral setae (first presutural dorsocentral barely distinguishable from general setulae); acrostichal setae not differentiated. Fore tibia approx. 2 times as long as first protarsomere. Hind tibia with 3 dorsal preapical setae and 3–4 well-developed anterodorsal and posterodorsal setae. Costal sector cs2 setulose ventrally. Abdomen: tergite 5 approx. as long as tergite 4. Male terminalia: surstylus triangular in lateral view; cerci stout with a slight restriction at approx. mid length; apical 1/5 of cerci not pointed. Female. Female differs from male as follows: Head: fronto-orbital plate entirely shiny and parafacial with less dense microtomentum. Frontal vitta approx. 0.2 times as wide as fronto-orbital plate in female. Parafacial 1.2 times as wide as postpedicel. Frons approx. 0.9 times as wide as compound eye in dorsal view.

Distribution

Neotropical – Trinidad and Tobago (Trinidad).

Etymology

The species epithet is derived from the Spanish word for Caribbean and should be treated as a Latin adjective.

Oriental region

Kinabalumyia Cerretti & Pape, gen. nov.

Fig. 18

Unambiguous character state changes

(Table 1, Fig. 20). Global apomorphies: 81:1; local apomorphies: 21:1, 53:1, 59:1.

Diagnosis

Head : head higher than long in lateral view. Facial ridge approx. 1.3 times as long as frons. Ocellar setae present though small. Frons 0.9–1.1 times as wide as compound eye in dorsal view. Median vertical setae strong and slightly converging. Two or three slightly convergent and reclinate frontal setae, descending to approx. half level of pedicel. Fronto-orbital plate nearly bare with one proclinate orbital seta (no sexual dimorphism). One upper lateroclinate orbital seta. Parafacial bare, at its narrowest point approx. twice as wide as maximum diameter of arista. Vibrissal angle strongly receding. Vibrissa well developed, arising slightly below the level of lower facial margin. Lower facial margin not sunken and slightly visible in lateral view. Facial ridge strongly concave with two or three short setulae above vibrissa. Face concave but antennae not hidden and clearly visible in lateral view. Antenna long and wide (in lateral view), much longer than height of gena. Postpedicel 4.5–6.5 times as long as pedicel. Postpedicel axe-shaped in male, more or less stick-like in female. Arista bare (or apparently so). First and second aristomere slightly thickened and strongly elongated. Lunule bare. Gena, in profile, approx. 3/5 (male), 1/2 (female) as high as compound eye. Palpus pretty short, dark brown.

Thorax : prosternum bare. Postpronotum with two setae. Two postsutural supra-alar setae (first postsutural supra-alar seta absent). Scutellum with one pair of strong lateral setae and one pair of short, crossed and horizontal apical setae. Anatergite with a tuft of short setulae below lower calypter. Subscutellum moderately swollen, mostly, though not entirely, sclerotised. Metathoracic spiracular lappets nearly undeveloped. Lower calypter distinctly tongue-shaped (ground-plan trait of Rhinophoridae) (Fig. 2E). Costal sector cs2 setose ventrally. Costal spine not differentiated from general costal setae. Costal sector cs5 clearly longer than costal sector cs2. Vein R1 entirely bare. Vein R4+5 bare dorsally. Bend of vein M1 absent. Crossvein dm-m forming a right angle with proximal section of M4. Vein CuA+CuP reaching wing margin. Preapical anterodorsal seta of fore tibia slightly longer than preapical dorsal seta. Fore tarsus not compressed. Tibiae of mid and hind leg normally developed. Hind tibia with three dorsal preapical setae.

Abdomen : tergites virtually without microtomentum and with relatively strong and recumbent general setulae; syntergite 1+2, tergites 3 and 4 with a pair of strong median marginal setae, without median discal setae, tergite 5 with strong marginal setae without discal setae.

Male terminalia : posterior margin of sternite 5 slightly concave, almost straight. Tergite 6 plate-like, with median marginal setae; tergite 6 and syntergosternite 7+8 fused. Connection between sternite 6 and syntergosternite 7+8 fused on right side. Cerci well developed and entirely fused medially into a syncercus. Surstylus long, narrow (almost stick-like), slightly enlarged distally and gently curved anteriorly. Surstylus not fused to epandrium. Bacilliform sclerite articulated (i.e., not fused) to laterobasal margin of surstylus. Hypandrial arms not fused medially. Connection between phallic guide and pregonite membranous. Pregonite well developed and lobe-like. Postgonite without anterior seta. Epiphallus well developed apically, pointed and attached dorsomedially to basiphallus. Extension of dorsal sclerite of distiphallus divided medially into two hemisclerites which are proximally fused to dorsal sclerite of distiphallus. Median process of ventral sclerotisation of distiphallus present, divided longitudinally and interrupted proximally, i.e., not connected to ventral plate. Acrophallus simple and scale-like spinules not differentiated.

Distribution

Oriental – Indonesia (Bali), Malaysia (Sabah), Philippines (Palawan).

Type species

Kinabalumyia pinax Cerretti & Pape, sp. nov., by present designation.

Etymology

The generic name is a composite word formed from the name of the type locality of the type species, Mount Kinabalu, and from the Greek word μύγα (miga), meaning fly. The name should be treated as a feminine noun.

Figure 18. 

Kinabalumyia gen. nov. A–C, F–I Kinabalumyia pinax sp. nov. (Malaysia, Sabah) D–E Kinabalumyia sp. ♀ (Philippines) A K. pinax sp. nov., habitus in lateral view B K. pinax sp. nov., abdomen in dorsal view C K. pinax sp. nov., head in lateral view D Kinabalumyia sp. ♀, head in frontal view, ♀ E Kinabalumyia sp. ♀, head in lateral view F K. pinax sp. nov., wing G–I K. pinax sp. nov.: epandrial complex in posterior (G) and lateral (H) view I hypandrial complex in lateral view [A–C holotype; F–I paratypes].

Kinabalumyia pinax Cerretti & Pape, sp. nov.

Fig. 18A–C, F–I

Type material

Holotype ♂: At light //SABAH: Mt Kinabalu /Mesilau 14.II.1964. J. Smart. /Royal Soc. Exped. /B.M. 1964-250 (NHMUK). Paratype ♂: same data and repository as holotype.

Description

Male. Body length: 3.5–3.7 mm. Colouration: fronto-orbital plate and upper occiput blackish-brown, frontal vitta black, remainder of head yellow; head covered with silver reflecting microtomentum. Scape and pedicel brown, postpedicel blackish-brown, palpus brown; postpronotum, proepisternum, proepimeron, and prosternum pale yellow, scutum, notopleuron, scutellum and subscutellum brown; anepisternum and upper anterior part of katepisternum brown, remaining sclerites pale brown except anatergite which is dark brown; tegula brown, basicosta and wing veins yellow; coxae and trochanters pale yellow, femora, tibiae and tarsi blackish-brown; abdomen can vary from entirely black to mostly black except yellow along posterior margin, on sides of syntergite 1+2 and along anterior margin of tergite 3; microtomentum virtually absent. Wing membrane hyaline. Head: frontal vitta well developed and slightly wider than fronto-orbital plate (both measured at midlength). Parafacial entirely bare below lower frontal seta. Lateral vertical seta not differentiated from strongest uppermost postocular setae. Prementum stout, not longer than wide; labella broad. Thorax: one posthumeral seta (medial); 1 + 1 supra-alar setae; 0 + 1–2 intra-alar setae; 2(3) + 3 dorsocentral setae; acrostichal setae not differentiated. One or two katepisternal setae. Legs: fore tibia without posterior seta. Abdomen: mid-dorsal depression on syntergite 1+2 confined to anterior half.

Distribution

Oriental – Malaysia (Sabah).

Etymology

The species epithet, which should be treated as a noun in apposition, is derived from the Greek noun pinax, meaning painting, in reference to the remarkable colour pattern of the thorax and abdomen.

Kinabalumyia sp. 1 (unidentified)

Fig. 18D, E

Material examined

1 ♀: Philippines, Palawan /Mantalingajan /Tagembung 1150 meter /18 Sept. 1961 /Noona Dan Exp. 61-62 //Caught by /Mercury-light /18.00–06.00 (NHMD).

Remarks

This female specimen, in fair condition, strongly resembles the two males from Sabah described as K. pinax above and may be conspecific with them. However, the sparse material available, the lack of females from the type locality, and the wide geographic separation provide sufficient taxonomic uncertainty to suggest caution. We therefore prefer to treat this specimen as unidentified awaiting further material.

Distribution

Oriental – Philippines (Palawan).

Kinabalumyia sp. 2 (undescribed)

Material examined

1 ♂: Indonesia, Bali, Bratan L. env., 1250m, Febr, 2014, O. Kosterin (ZMUM); photo only (Fig. 19), image also available from www.diptera.info (photo_id=9408).

Remarks

The photo shows a male specimen assessed as belonging to the genus here newly described as Kinabalumyia. Shape of the antenna and colouration of the body suggest that it is not conspecific with K. pinax and therefore represents an undescribed species.

Distribution

Oriental – Indonesia (Bali).

Figure 19. 

Kinabalumyia sp. 2, undescribed species from Indonesia, Bali (ZMUM).

Diversity and distribution

The Palaearctic Region hosts by far the most diverse rhinophorid fauna, with 92 of the 177 described species, and these are heavily concentrated to the western part. The Afrotropical Region contains 33 species, followed by the Australasian and Neotropical regions with 24 and 21 species, respectively. Only seven species (plus the two undescribed) have been recorded from the Oriental Region, and the by far most species-poor region is the Nearctic with only four native rhinophorids, all belonging to the genus Bezzimyia. This genus reaches its peak of diversity in the Neotropical Region. Indeed, the distribution of the Nearctic species of Bezzimyia is limited to the southernmost parts of the USA (Arizona, Texas, Georgia, Florida), corresponding with the range of the few native Nearctic woodlice (see Jass and Klausmeier 2000, Schmalfuss 2003), which are the most probable hosts of Bezzimyia and relatives. The near-absence of native woodlice in most of the temperate parts of North America has been given as in need of explanation (Jass and Klausmeier 2000), although it is rather the remarkable diversity of woodlice in the western Palaearctic that asks for scrutiny (“Paradoxically, their present centre of distribution and evolution is the region of the Mediterranean Sea”, Hatch 1947: 175; see also Sfenthourakis and Hornung 2018).

Faunistic connections are scarce among biogeographic regions, as virtually no rhinophorid species is so far known to be distributed across two or more regions, with the exclusion of a few anthropogenic introductions outside their native distributional range. In particular, three rhinophorids have been introduced from Europe to the Americas (Downes 1965, Mulieri et al. 2010, O’Hara et al. 2015, Wood et al. 2018), namely: Melanophora roralis (Linnaeus), which is now widespread in the New World; Stevenia deceptoria (Loew), recently reported from Ohio (USA) and northern Argentina; and Phyto discrepans Pandellé, known from Canada. While the two former species are now well established, little is known about the status of the latter. Moreover, Melanophora roralis, which is a native West-Palaearctic element, has been introduced to Japan and Cape Verde Islands, where it is now established (Crosskey 1977, Kato and Tachi 2016).

Phylogeny and suprageneric classification

Monophyly of Rhinophoridae and differences with previous analyses

As stressed by Cerretti and Pape (2012) and Cerretti et al. (2014a), the rhinophorid phylogeny based on currently available morphological data is poorly resolved, while a comprehensive phylogenetic study based on molecular data is still under completion (Gisondi et al., unpublished). Nevertheless, refined morphological evidence, coupled with new discoveries, are producing interesting results. Our taxon sample includes at least one species of all recognised rhinophorid genera, thereby comprising 14 more ingroup species than that of Cerretti et al. (2014a) (Table 1, asterisked names).

Our analyses reconstructed Rhinophoridae (clade A) as monophyletic based on eight unambiguous character state changes [four traceable in the adult: arista with short trichia (1:0, local apomorphy), first postsutural supra-alar seta short (25:1, local apomorphy), subscutellum moderately swollen (26:1, global apomorphy), lower calypter tongue-shaped (36:1, local apomorphy); four in the first instar larva: antenna long and tapering (85:1, global apomorphy), posterior part of anal division modified as a terminal sucker (86:1, global apomorphy), mandibles toothed or serrated (88:1, global apomorphy), and parastomal bar of cephaloskeleton long and slender (91:1, local apomorphy)], confirming the split of the family into two main subclades: the Phyto group (clade B), the first instar larvae of which move by somersaulting, and the Stevenia group (clade E), whose first instars move in a leech-like crawling fashion (see Bedding 1973, Pape 1986, Pape and Arnaud 2001, Cerretti et al. 2014a) (Fig. 20). Analyses by Cerretti et al. (2014a) found the New Caledonian endemic genus Rhinodonia as sister to the remaining rhinophorids, and the Australasian genera Axinia and Bixinia as sister taxa nested within the Stevenia group. Our analyses still recovered Bixinia within the Stevenia group but notably diverged from Cerretti et al. (2014a) in retrieving Rhinodonia and Axinia as sister clades and part of an Oriental–Australasian radiation of the Phyto group. The differences between analyses stand likely by the double effect given by the new taxa included in the present version of the matrix and by the application of the implied weighting function implemented in TNT [k/(es+k), where k = concavity constant and es=extra step, i.e., as k increases, the function approximate the linear, equally weighted (“unweighted”) function (Goloboff 1993)]. Under implied weighting, the increase in fit (with differing strength depending upon the k-value) caused by the reduction in number of steps for a character is higher for characters with low homoplasy because each step represents a larger fraction of the total homoplasy for that character. In Fig. 20 one of these apomorphies, shared by the Australasian taxa except Bixinia (clade C), is the lack of a bend of vein M1 (45:1). The position of Axinia as a member of the Phyto group has recently been corroborated by molecular data (Cerretti et al. 2019). However, the affinities of Bixinia are more problematic, as both its separation from Axinia and its position within a Neotropical clade (I) appear unlikely, as ‘intuitively’ assessed by its relative similarity to other Australasian taxa (see below).

Figure 20. 

Phylogeny of Rhinophoridae based on morphological evidence. Single tree obtained under IW (total fit: 62.33), enforcing a k-value of 4, with unambiguous character state changes mapped above branches [black squares: global apomorphies (i.e., uncontradicted and unreversed apomorphic character states; white squares: local apomorphies (i.e., homoplasious character state due to convergence or reversal)] and new taxa highlighted in bold.

Phyto group and Stevenia group

The Phyto group (Fig. 20, clade B) was supported by four unambiguous character state changes [anal division of the first instar larva with longitudinal cuticular ridges in posteroventral position (95:1, global apomorphy) and a tongue-like projection in posterodorsal position (96:1, global apomorphy), cephaloskeleton of the first instar with longitudinal incision on parastomal bar (92:1, local apomorphy), and adult male with well-developed proclinate orbital setae (9:1, local apomorphy)]. The clade is composed exclusively of Old World elements. Remarkably, most of the characters whose derived condition supported the monophyly of clade B (Phyto group) are from the first instar larva, which is known for only three (i.e., Paykullia, Melanophora, and Phyto) of the 12 genera composing the group.

The Stevenia group (clade E) is nearly worldwide in distribution. Its monophyly was supported by four unambiguous character state changes [first instar larva with prolegs (93:1, global apomorphy), mandible provided with three or more teeth (99:0, global apomorphy), anal division with pair of vesicles in posteroventral position (97:1, local apomorphy), and adult male with bacilliform sclerite firmly fused to laterobasal margin of surstylus (65:1, local apomorphy)]. This clade was in turn divided into three subclades: F, H and I. Clade F is a relatively large assemblage supported by one global apomorphy [first instar larva with dorsoventrally flattened body (90:1)], and composed by Old World taxa widespread in the Afrotropical (Maurhinophora gen. nov., Melanomyiodes, Queximyia, Stevenia [in part]), Oriental (Acompomintho [in part], Stevenia [in part]), and Palaearctic (Acompomintho [in part], Azaisia, Macrotarsina, Baniassa, Metoplisa, Oplisa, Rhinomorinia, Rhinophora, Tricogena, Tromodesia, Stevenia [in part]) regions. Clade H was supported by two unambiguous character state changes of adult male [dorsal sclerite of acrophallus semicylindrical in shape (80:1, global apomorphy), and parafrontal plate with proclinate orbital setae (9:1, local apomorphy)] and includes only the endemic Afrotropical genus Ventrops (Cerretti & Pape, 2012). Finally, clade I is weakly supported by three local apomorphies [female without proclinate orbital setae (10:0), male surstylus fused to epandrium (66:1), and postgonite without anterior seta (69:1)], and is mostly composed of Neotropical taxa, with the notable exception of the Australian genus Bixinia, whose phylogenetic position seems unlikely and needs further study. Ventrops (i.e., clade H) was reconstructed as sister to clade I based on two local apomorphies [fronto-orbital plate with more than two proclinate orbital setae (11:1), and extensions of dorsal sclerite of distiphallus entirely fused mid-dorsally from each other into a single sclerotisation (71:1)]. The preimaginal stages and the natural history of Ventrops are unknown and more data are needed to clarify its phylogenetic position within Rhinophoridae.

The recent molecular-based phylogenetic reconstruction by Cerretti et al. (2019) recovered monophyly for both the Phyto group and the Stevenia group.

The Aporeomyia conundrum

The genus Aporeomyia was originally assigned to the family Tachinidae by Pape and Shima (1993), despite sharing with Rhinophoridae two local apomorphies, i.e., a tongue-like lower calypter (36:1) and anterior and posterior lappets of metathoracic spiracle subequal in size and directed outward (30:1). The original placement of this unusual taxon was based on the presence of a moderately convex subscutellum and a distiphallus characterised by an undivided extension of the dorsal sclerite, as in tachinids (Pape and Shima 1993). Aporeomyia is here reassigned to Rhinophoridae based on a reassessment of the homologies of the thorax and male terminalia. The subscutellum in Aporeomyia is moderately convex, but not bulging and fully sclerotised as in tachinids, which indicates a rhinophorid affinity. Notably, several rhinophorids also have a moderately convex subscutellum similar to that of Aporeomyia, in particular among species within various genera of the Phyto group, such as Phyto, Comoromyia, Malayia, Rhinopeza, and Kinabalumyia gen. nov. This character state is not restricted to this group: the Australasian genus Bixinia has a bulging subscutellum similar to that of tachinids, but its inclusion in Rhinophoridae is beyond any reasonable doubt (Kutty et al. 2019). As discussed by Pape and Shima (1993: 78, fig. 5), the extension of the dorsal sclerite of the distiphallus in Aporeomyia antennalis Pape & Shima is undivided, as typical for nearly all tachinids and a few other non-tachinid oestroids (see also Rognes 1997, Pape and Arnaud 2001). However, recent morphology-based cladistic analyses including a growing number of rhinophorid taxa indicate that an undivided extension of the dorsal sclerite of the distiphallus characterises several rhinophorid species belonging to different genera [e.g., Rhinophora lepida (Meigen), some Bezzimyia, Shannoniella, Trypetidomima, some Melanophora], thus confirming this trait as homoplasious within oestroids, and among rhinophorids in particular (Pape and Arnaud 2001, Cerretti and Pape 2012, Cerretti et al. 2014a, 2017). Moreover, the apparently undescribed species of Aporeomyia from Sabah included in our analysis is characterised by an extension of the dorsal sclerite of distiphallus that is longitudinally divided, even if shallowly, into two branches, suggesting that the two conditions can occur even in closely related species. We retrieved Aporeomyia nested within the Phyto group (clade B), as sister to the Oriental genus Kinabalumyia gen. nov. based on three unambiguous character state changes [first aristomere at least four times as long as wide (6:2, global apomorphy), wing vein CuA+CuP reaching wing margin (50:1, local apomorphy), and male with connection between tergite 6 and syntergosternite 7+8 fused (58:1, global apomorphy)]. The monophyly of Aporeomyia is based almost entirely on the strongly autapomorphic postpedicel with a tripartition (Fig. 6B) similar to that of the Neotropical Bezzimyia lobata Pape & Arnaud, but with sufficient differences to be considered a unique autapomorphy of Aporeomyia. In conclusion, we found strong evidence supporting Aporeomyia as a member of the Rhinophoridae as well as generic monophyly.

The position of Kinabalumyia gen. nov. within the Phyto group

The Oriental genus Kinabalumyia gen. nov. was supported by four unambiguous character state changes [distiphallus with a helmet-shaped, partly sclerotised envelope (81:1, global apomorphy), palpus reduced (21:1, local apomorphy), and male with: sternite 5 without posterior notch (53:1, local apomorphy), and connection between sternite 6 and syntergosternite 7+8 fused on right side (59:1, local apomorphy)] and it was reconstructed as sister to the Oriental genus Aporeomyia (see above). This clade was in turn retrieved as sister to the Australasian clade D (Rhinodonia + Axinia) based on two local apomorphies [base of wing vein R4+5 bare (43:1), and male cerci medially fused into a syncercus (61:1)]. We think that the helmet-shaped envelope of distiphallus is a strong autapomorphy supporting the erection of a new genus for the examined specimens. We have examined four specimens of Kinabalumyia gen. nov. from three localities scattered from Bali in Indonesia to Palawan in the Philippines, suggesting that this genus is widespread at least throughout the non-continental part of the Oriental Region, with further species probably awaiting discovery. Clade C (Fig. 20) was supported by two unambiguous character state changes [bend of vein M1 indistinct (45:1, global apomorphy) and metathoracic spiracular lappets absent (30:0, local apomorphy)] and it reconstructed the Australasian clade (clade D) arising from a grade composed of Australasian (Rhinopeza) and Oriental (Aporeomyia, Kinabalumyia gen. nov.) taxa. Despite being weakly supported by one local apomorphy [arista thickened at least on basal 3/4 (5:1)], clade C is in turn sister to the Oriental genus Malayia. If this reconstruction were accurate, the hypothesis of Cerretti et al. (2014a) about the origin of the Australasian rhinophorid fauna would need to be reconsidered. In fact, our phylogeny favours an Old World ancestor for most of the Australasian taxa, the only notable exception being Bixinia of Neotropical descent.

The Stevenia group and the position of Maurhinophora gen. nov., Marshallicona gen. nov. and Neotarsina gen. nov.

The monotypic genus Maurhinophora gen. nov. was erected to accommodate the new species M. indoceanica sp. nov. from Mauritius, based on a single female. The genus is supported in this work by six local apomorphies [facial plate and lower facial margin deeply sunken (13:1 and 14:1), palpus absent (21:2), lappets of metathoracic spiracle unequal in size, i.e., posterior one distinctly larger (30:2), wing vein R1 with a row of setulae along whole length (42:1), and base of wing vein R4+5 with a row of setae reaching at least crossvein r-m (44:1)] and was reconstructed as sister to the monotypic endemic Afrotropical genus Queximyia. Maurhinophora gen. nov. and Queximyia were part of the large clade F. The latter group was based on a global apomorphy of the first instar larva [first instar larva with slightly flattened body shape (90:1)], while an included subclade (clade G), also comprising Maurhinophora gen. nov. + Queximyia, was instead supported by one global apomorphy of the male terminalia [acrophallus distinctly tripartite, i.e., with three openings (79:1)]. This phylogenentic reconstruction suggests that the first instar and adult male of Maurhinophora gen. nov. (both unknown) might share these traits as well.

Two of the four newly described genera are Neotropical in distribution. During the last decades, the native New World rhinophorid fauna has increased remarkably from one genus and two species to the present six genera and 25 species (Pape and Arnaud 2001, Pape 2010, Cerretti et al. 2014a, Nihei and Andrade 2014, Nihei et al. 2016), and more are most likely awaiting discovery. Although based on a large array of local apomorphies, our analysis corroborated the non-monophyly of the genus Bezzimyia, as previously suggested by Cerretti and Pape (2012) and Cerretti et al. (2014a). The monotypic genus Marshallicona gen. nov. was supported by seven local apomorphies [arista microtrichia bottlebrush-like in male (2:1), male with proclinate orbital setae (9:1), base of wing vein R4+5 bare (43:1), abdominal tergites 3 and 4 without median marginal setae (51:1 and 52:1), abdominal tergite 6 bare in male (56:1), connection between surstylus and epandrium membranous (66:0), and phallic guide fused to pregonites (68:1)] and was reconstructed as sister to the clade Trypetidomima + Shannoniella.

The genus Neotarsina gen. nov. was erected for two new species from Peru (N. andina sp. nov.) and Trinidad and Tobago (N. caraibica sp. nov.). As described above for Kinabalumyia gen. nov., we suspect that Neotarsina gen. nov. is widespread in the Neotropics and also more diverse, not just including the presently described species. Neotarsina gen. nov. was reconstructed as sister to Bezzimyia (B. barbarista Pape & Arnaud + B. busckii Townsend) based on three unambiguous character changes [female with a groove between fronto-orbital plate and parafacial (8:1, global apomorphy), male fore tarsus laterally compressed (31:1, local apomorphy), and bacilliform sclerite articulated (i.e., not fused) to laterobasal margin of surstylus (65:0, local apomorphy)], and its monophyly relies on four unambiguous character changes [tibia of mid and hind legs laterally compressed and distinctly keeled dorsally (33:1, global apomorphy), palpus reduced (21:1, local apomorphy), male with connection between sternite 6 and syntergosternite 7+8 on right side (59:1, local apomorphy), and connection between dorsal sclerite of distiphallus and extensions of dorsal sclerite of distiphallus sclerotised (72:1, local apomorphy)].

Catalogue

Family Rhinophoridae Robineau-Desvoidy, 1863

Rhinophoridae Robineau-Desvoidy 1863: 3. Type genus: Rhinophora Robineau-Desvoidy 1830.

Genus Acompomintho Villeneuve, 1927

Acompomintho Villeneuve 1927: 223. Type species: Acompomintho lobata Villeneuve, 1927, by monotypy.

Wagneriopsis Townsend 1927: 281. Type species: Wagneriopsis formosensis Townsend, 1927, by original designation.

[Acampomintho]: Townsend (1931: 383), Lopes (1938: 555), Herting (1961: 5); incorrect subsequent spelling of Acompomintho Villeneuve, 1927.

caucasica (Villeneuve, 1908), stat. rev., comb. nov.

Frauenfeldia caucasica Villeneuve 1908: 287. Type locality: Russia, Krasnodar Krai, Tuapse. Holotype male not located [“coll. Schnabl”; “C. Tuapse”].

Remarks. Treated as unplaced to genus by Herting (1961: 34) but here resurrected as a valid species in the genus Acompomintho. Herting (1961) did not examine the holotype of F. caucasica and he claimed that the original description was inconclusive as to the generic placement of caucasica. In particular, Herting mentioned that despite similarities in colouration and habitus with Tricogena rubricosa (Meigen) [Frauenfeldia Egger is a junior synonym of Tricogena, see below], the original description of F. caucasica lacked any mention of robust setae on the lower half of the parafacial (characterising Tricogena and Stevenia, but not all described Acompomintho), or any information about other diagnostic characters. We have not been able to locate and examine the holotype of caucasica, but we consider the original description to match Acompomintho better than Tricogena.

Distribution. Palaearctic – Russia (North Caucasus).

itoshimensis Kato & Tachi, 2016

Acompomintho itoshimensis Kato and Tachi 2016: 83. Type locality: Japan, Kyushu, Fukuoka, Shimasakurai. Holotype male in BLKU.

Distribution. Palaearctic – Japan (Kyushu).

lobata Villeneuve, 1927

Acompomintho lobata Villeneuve 1927: 223. Type locality: Taiwan. Syntypes, male(s) and female(s), in DEI, NHMUK, USNM, possibly also elsewhere.

Wagneriopsis formosensis Townsend 1927: 282. Type locality: Taiwan: Hokuto, Anping, Kankau and Macuyama. Syntypes, male(s) and female(s), 1 male and 6 females in DEI, possibly also elsewhere.

Distribution. Palaearctic – North Korea, South Korea. Oriental – Japan (Ryukyu Is), Taiwan.

sinensis (Villeneuve, 1936), stat. rev., comb. nov.

Frauenfeldia sinensis Villeneuve 1936: 7. Type locality: China, southern Gansu. Holotype female not located.

Remarks. Treated as unplaced to genus by Herting (1961: 34) but here resurrected as a valid species in the genus Acompomintho. The revised status of this nominal species is based on the recent discovery in Tajikistan of two female specimens (Table 4) belonging to the genus Acompomintho, which match the original description of F. sinensis.

Distribution. Palaearctic – China (Gansu), Tajikistan (Gorno-Badachšan, Rŭshan) [new record].

Genus Apomorphyto Cerretti, Lo Giudice & Pape, 2014

Apomorphyto Cerretti et al. 2014a: 675. Type species: Apomorphyto inbio Cerretti, Lo Giudice & Pape, 2014, by original designation.

inbio Cerretti, Lo Giudice & Pape, 2014

Apomorphyto inbio Cerretti et al. 2014a: 676. Type locality: Costa Rica, Guanacaste, Estación Las Pailas, Parc Nacional Rincón de la Vieja. Holotype male in INBio.

Distribution. Neotropical – Costa Rica, Nicaragua.

Genus Aporeomyia Pape & Shima, 1993

Aporeomyia Pape and Shima 1993: 77. Type species: Aporeomyia antennalis Pape & Shima, 1993, by original designation.

antennalis Pape & Shima, 1993

Aporeomyia antennalis Pape and Shima 1993: 77. Type locality: Philippines, Mindanao, Mt. Apo, Lake Agco. Holotype male in BLKU.

Distribution. Oriental – Philippines (Mindanao).

Undescribed sp.: Malaysia (Sabah) (CNC, NHMD).

Genus Axinia Colless, 1994

Axinia Colless 1994a: 484. Type species: Axinia arenaria Colless, 1994, by original designation.

Remarks. There are two spellings of this genus-group name in Colless (1994): Axinia on page 484 plus another 90 occurrences, and Axinis on page 502, used in the combination “Axinis cornuta”. We consider the multiple use of the spelling “A. cornuta” in contexts where the “A.” is evidently an abbreviation for “Axinia”, as “clear evidence of an inadvertent error” (Code, Article 32.5.1), and Axinis must accordingly be corrected to Axinia.

Dixicera Colless 1994a: 508 (as subgenus of Axinia Colless, 1994). Type species: Axinia carnei Colless, 1994, by original designation.

Barrinea Colless 1994a: 511. Type species: Barrinea disjuncta Colless, 1994, by original designation.

Ismaya Colless 1994a: 512. Type species: Ismaya miranda Colless, 1994, by original designation. [Junior homonym of Ismaya Bouček, 1988 (Hymenoptera: Chalcidoidea).]

Chirops Colless 1994a: 516. Type species: Chirops arcana Colless, 1994, by original designation.

Johnismaya Colless 1994b: 380. New name for Ismaya Colless, 1994.

[Axinis]: Colless (1994a: 502); incorrect original spelling of Axinia Colless [teste this work].

arcana (Colless, 1994)

Chirops arcana Colless 1994a: 517. Type locality: Papua New Guinea, E end Saruwaged Range, 20 km SSW Kabwum, 2550 m. Holotype male in BPBM.

Distribution. Australasian – Papua New Guinea (PNG).

arenaria Colless, 1994

Axinia (Axinia) arenaria Colless 1994a: 486. Type locality: Australia, Western Australia, 37 mi N Ajana. Holotype male in ANIC.

Distribution. Australasian – Australia (Northern Territory, South Australia, Western Australia).

austrina Colless, 1994

Axinia (Axinia) austrina Colless 1994a: 496. Type locality: Australia, Australian Capital Territory, Blundells Creek. Holotype male in ANIC.

Distribution. Australasian – Australia (ACT, New South Wales).

bicolor Colless, 1994

Axinia (Axinia) bicolor Colless 1994a: 505. Type locality: Australia, Queensland, Lamington National Park, O’Reilly’s. Holotype male in ANIC.

Distribution. Australasian – Australia (Queensland).

brevicentrum Colless, 1994

Axinia (Axinia) brevicentrum Colless 1994a: 503. Type locality: Australia, Western Australia, 37 km SW Mt Ragged. Holotype male in ANIC.

Distribution. Australasian – Australian (South Australia, Victoria, Western Australia).

cantrelli Colless, 1994

Axinia (Axinia) cantrelli Colless 1994a: 489. Type locality: Australia, Queensland, Mt Tamborine. Holotype male in QDPC.

Distribution. Australasian – Australia (New South Wales, Queensland).

carnei Colless, 1994

Axinia (Dixicera) carnei Colless 1994a: 508. Type locality: Australia, Western Australia, 8 mi ENE Millstream. Holotype male in ANIC.

Distribution. Australasian – Australia (Western Australia).

cornuta Colless, 1994

Axinia (Axinia) cornuta Colless 1994a: 502. Type locality: Australia, Australian Capital Territory, Blundells Creek. Holotype male in ANIC.

Distribution. Australasian – Australia (ACT, Tasmania, Victoria).

disjuncta (Colless, 1994)

Barrinea disjuncta Colless 1994a: 511. Type locality: Australia, Queensland, Lake Barrine. Holotype male in ANIC.

Distribution. Australasian – Australia (Queensland).

gressitti Colless, 1994

Axinia (Axinia) gressitti Colless 1994a: 501. Type locality: Papua New Guinea, Morobe Province, Mt Missim, 1600 m. Holotype male in BPBM.

Distribution. Australasian – Papua New Guinea (Morobe).

lucaris Colless, 1994

Axinia (Axinia) lucaris Colless 1994a: 492. Type locality: Australia, Queensland, 13 km NE by N Yungaburra, Danbulla Forest Reserve. Holotype male in ANIC.

Distribution. Australasian – Australia (New South Wales, Queensland).

minuta Colless, 1994

Axinia (Dixicera) minuta Colless 1994a: 509. Type locality: Australia, Western Australia, Crowea near Busselton. Holotype male in ANIC.

Distribution. Australasian – Australia (Western Australia).

miranda (Colless, 1994)

Ismaya miranda Colless 1994a: 514. Type locality: Papua New Guinea, Northern Province, Mt Scratchley, 3500 m. Holotype male in ANIC.

Distribution. Australasian – Papua New Guinea (Northern Province).

mutabilis Colless, 1994

Axinia (Axinia) mutabilis Colless 1994a: 499. Type locality: Australia, Queensland, Lamington National Park, O’Reilly’s. Holotype male in ANIC.

Distribution. Australasian – Australia (Queensland).

naumanni Colless, 1994

Axinia (Axinia) naumanni Colless 1994a: 498. Type locality: Australia, Tasmania, 4 km E Rosebery. Holotype male in ANIC.

Distribution. Australasian – Australia (Tasmania).

zentae Colless, 1994

Axinia (Axinia) zentae Colless 1994a: 495. Type locality: Australia, Australian Capital Territory, Blundells Creek. Holotype male in ANIC.

Distribution. Australasian – Australia (ACT, New South Wales).

Genus Azaisia Villeneuve, 1939

Azaisia Villeneuve 1939: 350. Type species: Azaisia setitarsis Villeneuve, 1939, by original designation.

Azaisiella Villeneuve 1939: 351 (as subgenus of Azaisia Villeneuve, 1939). Type species: Azaisiella obscura Villeneuve, 1939, by original designation.

obscura Villeneuve, 1939

Azaisia (Azaisiella) obscura Villeneuve 1939: 352. Type locality: Madeira. Holotype female in IRSNB.

Distribution. Palaearctic – Azores?, Madeira.

setitarsis Villeneuve, 1939

Azaisia setitarsis Villeneuve 1939: 351. Type locality: Madeira. Holotype female in IRSNB.

Distribution. Palaearctic – Madeira.

Genus Baniassa Kugler, 1978

Baniassa Kugler 1978: 73. Type species: Baniassa fascipennis Kugler, 1978, by orginal designation.

fascipennis Kugler, 1978

Baniassa fascipennis Kugler 1978: 74. Type locality: Israel, Mt Carmel. Holotype male in TAU.

Distribution. Palaearctic – Israel.

fenestrata Zeegers, 2008

Baniassa fenestrata Zeegers 2008: 733, 734. Type locality: United Arab Emirates, Fujairah. Holotype male in NBCL.

Distribution. Palaearctic/Afrotropical – Oman, United Arab Emirates.

paucipila Pape, 1985

Baniassa paucipila Pape 1985: 209. Type locality: Iraq, Kurdistan, Arbil. Holotype male in USNM (unique specimen identifier: USNMENT01519730).

Distribution. Palaearctic – Iraq.

Genus Bezzimyia Townsend, 1919

Bezzimyia Townsend 1919a: 591. Type species: Bezzimyia busckii Townsend, 1919, by original designation.

Lutzomyia Curran 1934: 387. Type species: Lutzomyia americana Curran, 1934, by monotypy. [Preoccupied by Lutzomyia França, 1927 (Diptera: Psychodidae).]

Pseudolutzomyia Rapp 1945: 278. New name for Lutzomyia Curran, 1934.

[Pseudolutozmyia]: Rapp (1945: 278); incorrect original spelling of Pseudolutzomyia [teste Pape and Arnaud (2001: 261)].

Group A

bisecta Pape & Arnaud, 2001

Bezzimyia bisecta Pape and Arnaud 2001: 260, 265. Type locality: Costa Rica, San José, Braulio Carillo National Park, 10°07'N, 83°58'W, 1000 m. Holotype male in USNM (unique specimen identifier: USNMENT01519747).

Distribution. Neotropical – Costa Rica.

hansoni Pape & Arnaud, 2001

Bezzimyia hansoni Pape and Arnaud 2001: 260, 273. Type locality: Costa Rica, Cartago, La Cangreja, 1950 m. Holotype male in USNM (unique specimen identifier: USNMENT01519735).

Distribution. Neotropical – Costa Rica.

pittieri Pape & Arnaud, 2001

Bezzimyia pittieri Pape and Arnaud 2001: 261, 279. Type locality: Venezuela, Aragua, Henri Pittier National Park, Pico Periquito, 1680 m. Holotype male in MIZA.

Distribution. Neotropical – Venezuela.

ramicornis Pape & Arnaud, 2001

Bezzimyia ramicornis Pape and Arnaud 2001: 260, 281. Type locality: Ecuador, Pichincha, nr Tinalandia [hotel 16 km SE Santo Domingo], 1150 m. Holotype male in CNC.

Distribution. Neotropical – Ecuador.

yepezi Pape & Arnaud, 2001

Bezzimyia yepezi Pape and Arnaud 2001: 261, 286. Type locality: Venezuela, Aragua, Rancho Grande, 1100 m. Holotype male in MIZA.

Distribution. Neotropical – Venezuela.

Group B

americana (Curran, 1934)

Lutzomyia americana Curran 1934: 387. Type locality: USA, Arizona, Cobabi Mts., Sta. Cruz Village, 3100 ft. Lectotype male (by designation of Sabrosky 1963: 14) in AMNH.

[Lutzomyia latifrons]: Curran (1934: 396, 398); incorrect original spelling of americana Curran, 1934 [teste Sabrosky (1963: 14)].

Distribution. Nearctic – USA (Arizona).

barbarista Pape & Arnaud, 2001

Bezzimyia barbarista Pape and Arnaud 2001: 260, 263. Type locality: Costa Rica, Cartago, La Cangreja, 1950 m. Holotype male in USNM (unique specimen identifier: USNMENT01519746).

Distribution. Neotropical – Belize, Costa Rica.

bulbosa Pape & Arnaud, 2001

Bezzimyia bulbosa Pape and Arnaud 2001: 260, 267. Type locality: Mexico, Veracruz, Fortín de Las Flores, Sumidero, Planta de la Serveceria. Holotype male in