Synopsis of Cis Latreille (Coleoptera: Ciidae) from southern Africa

Abstract

A synopsis of the Cis Latreille, 1796 from southern Africa is provided, with the description of 10 new species: Cis bicaesariatus sp. n., Cis foveocephalus sp. n., Cis grobbelaarae sp. n., Cis lacinipennis sp. n., Cis makrosoma sp. n., Cis mpumalangaensis sp. n., Cis parvisetosus sp. n., Cis tessariplacus sp. n., Cis umlalaziensis sp. n. and Cis westerncapensis sp. n. The introduced species Cis fuscipes Mellié, 1849 is recorded for the first time from the Republic of South Africa. New geographic records are provided for the following species: Cis neserorum Souza-Gonçalves & Lopes-Andrade, 2017; Cis regius Orsetti & Lopes-Andrade, 2016 and Cis stalsi Souza-Gonçalves & Lopes-Andrade, 2017. Most southern African Cis are placed in available or newly proposed species-groups and a provisional identification key is provided.

1. Introduction

Ciidae is a cosmopolitan family and comprises of more than 700 described species in 51 genera. The genus Cis Latreille, 1796 (Ciinae: Ciini) has about 400 described species occurring in all biogeographic regions, except for the Antarctic [1,2,3]. It is the most diverse genus in the family, including more than half of all described Ciidae, but it is possibly polyphyletic [4,5]. The previously proposed division into subgenera is not in use and part of the Cis species is organized in artificial species-group [2,3,6,7].

The sub-Saharan Ciidae, most belonging to the Ethiopian region (biogeographic regions sensu Morrone [8]), are represented by 75 described species [1,9,10,11,12], of which 53 belong to Cis. In southern Africa, comprising Botswana, Lesotho, Namibia, Republic of South Africa, Swaziland and the southern tip of Mozambique, there are 19 described species of Cis: C. afer Fåhraeus, 1871, C. aster Souza-Gonçalves & Lopes-Andrade, 2017, C. bimucronatus Motschoulsky, 1851, C. caffer Fåhraeus, 1871, C. capensis Mellié, 1849, C. chinensis Lawrence, 1991, C. delagoensis Pic, 1916, C. makebae Souza-Gonçalves & Lopes-Andrade, 2017, C. mandelai Souza-Gonçalves & Lopes-Andrade, 2017, C. masekelai Souza-Gonçalves-& Lopes-Andrade, 2017, C. mooihoekite Souza-Gonçalves & Lopes-Andrade, 2018, C. muriceus Mellié, 1849, C. neserorum Souza-Gonçalves & Lopes-Andrade, 2017, C. paraliacus Souza-Gonçalves & Lopes-Andrade, 2018, C. pickeri Lopes-Andrade et al., 2009, C. regius Orsetti & Lopes-Andrade, 2016, C. stalsi Souza-Gonçalves & Lopes-Andrade, 2017, C. testaceus Fåhraeus, 1871 and C. urbanae Souza-Gonçalves & Lopes-Andrade, 2017.

The aim of this paper is to provide a synopsis of southern African Cis, with description of 10 new species. Most of them are placed in available or newly proposed species-groups. New geographic records for previously described species are provided, together with a provisional identification key to species occurring in southern Africa. The invasive Cis fuscipes Mellié, 1849 is recorded for the first time from the Republic of South Africa.

2. Materials and Methods

Museum abbreviations are as follows:

ANIC 

Australian National Insect Collection, CSIRO Entomology (Canberra, Australian Capital Territory, Australia)

CELC 

Coleção Entomológica do Laboratório de Sistemática e Biologia de Coleoptera da Universidade Federal de Viçosa (Viçosa, Minas Gerais, Brazil)

CUIC 

Cornell University Insect Collection (Ithaca, NY, USA)

SANC 

South African National Collection of Insects (Pretoria, Gauteng, Republic of South Africa)

The new species described here (Figures 1–2, Figures 4–11) and part of the new records of previously described species (Figure 3) are based on specimens collected and organized by the staff of the SANC, mostly during 10 years of a project on parasitoids of Ciidae in the Republic of South Africa [9]. The original coding of the morphospecies [9] has been revised and some of them have been described [10,11,12,13]. Correspondence of morphospecies and the new species are cited in the subheadings of each new species.

Terms for external morphology and male terminalia of ciids follow Lawrence [3], Lawrence et al. [14] and Lawrence & Lopes-Andrade [15,16], but see also Oliveira et al. [2] for an explanation on the use of “tegmen”. The following abbreviations are used for measurements (in mm) and ratios: BW (width of anterior edge of scutellar shield), CL (length of antennal club measured from base of the eighth to apex of the tenth antennomere), EL (elytral length along the midline), EW (greatest width of elytra), FL (length of antennal funicle measured from base of the third to apex of the seventh antennomere), GD (greatest depth of body measured in lateral view), GW (greatest diameter of eye), PL (pronotal length along midline), PW (greatest pronotal width), SL (length of scutellar shield), TL (total length counted as EL+PL, i.e., excluding head). The GD/EW and TL/EW ratios indicate the degree of body convexity and elongation, respectively.

Transcription of labels, dissection, photography and measurement of specimens follow the methods provided by Araujo & Lopes-Andrade [17]. Exemplar specimens of the new species from all localities were measured. The number of measured specimens depended on availability and observed variation in size. Differences are given in “Variation”, together with standard measurements (mean and standard deviation) and ratios. Data on host fungi extracted from labels and literature are summarized in the corresponding sections and names were updated consulting the database Index Fungorum [18], together with a corresponding number of records in each fungus species and indicating breeding records. The criteria provided in Orledge and Reynolds [19] was followed for determining breeding records. The distribution maps (Figures 13–15) were created in the freeware QGIS 2.14.2-Essen [20]. Throughout the identification key, provinces of the Republic of South Africa are indicated as EC (Eastern Cape), FS (Free State), GP (Gauteng), KZN (KwaZulu-Natal), LP (Limpopo), MP (Mpumalanga), NW (North West) and WC (Western Cape).

The following males were dissected: one Cis bicaesariatus sp. n. from Die Hel Nature Reserve, Mpumalanga Province; two Cis foveocephalus sp. n., from Mooihoek Farm, Mpumalanga Province; four Cis grobbelaarae sp. n. (three from Die Hel Nature Reserve, Mpumalanga Province; and one from D’Nyala Nature Reserve, Limpopo Province); three Cis lacinipennis sp. n. (two from Strathdene Farm and one from Cathedral Peaks Forest Station, KwaZulu-Natal Province); one Cis makrosoma sp. n. from Nelspruit, Mpumalanga Province; two Cis mpumalangaensis sp. n. from Mooihoek Farm, Mpumalanga Province; three Cis parvisetosus sp. n. (one from Monk’ S Cowl, one from Mpisini Nature Reserve, KwaZulu-Natal Province; and one from Prince Alfred’ S Pass, Western Cape Province); one Cis regius from Storms River Mouth Restcamp, Western Cape Province; two Cis tessariplacus sp. n. (one from Mooihoek Farm and one from Alkmaar, Mpumalanga Province); one Cis umlalaziensis sp. n. from Umlalazi Nature Reserve, KwaZulu-Natal Province; two Cis westerncapensis sp. n. (one from Montagu Pass and one from Saasveld Forestry College, Western Cape Province). The following females were dissected: three Cis fuscipes from Garden of Eden Indigenous Forest, Western Cape Province; two Cis mpumalangaensis sp. n. from Mooihoek Farm, Mpumalanga Province; one Cis westerncapensis sp. n. from Saasveld Forestry College, Western Cape Province. The sclerites of aedeagi shown in Figures 1(F–I) and 10(F–I) are of the holotypes. The sclerites of aedeagi shown in Figure 10(B–E, G–J) are of paratypes from other localities and demonstrate the low variation between populations. The sclerites of aedeagi shown in Figures 2(F–I), 4(F–I), 5(F–H), 6(F–H), 7(F–I), 8(F–I) and 11(E–G) are of paratypes from type localities. The sclerites of aedeagus shown in Figure 12(F–I) are of a paratype from a locality rather than the type locality, but identified as conspecific to those from the type locality. In the latter case, aedeagus extracted from male of the type locality was a little damaged during dissection and laid in a bad position on the slide. Information on the gula provided in the descriptions is restricted to the ratio of gula width to head width.

The identification key is restricted to species described and examined by us, because we did not have access to any identified material or type specimens of the other southern African species (C. afer, C. bimucronatus, C. caffer, C. capensis, C. delagoensis, C. muriceus and C. testaceus). The species are presented in alphabetical order.

3. Results

Information on C. afer, C. caffer, C. delagoensis, C. muriceus and C. testaceus is mostly that from the original descriptions [21,22,23], images of types of identified specimens made available to us (see Acknowledgments) and provided by Ferrer [24]. Information on C. bimucronatus and C. capensis are from the original descriptions [22,25]. The descriptions of C. delagoensis and C. bimucronatus are anecdotal; and only the holotype of C. muriceus is known, which is a female in poor condition that seems to be teneral (i.e., recently hatched, light-colored adult). Therefore, we have not considered C. bimocrunatus, C. delagoensis and C. muriceus in comparisons provided in the diagnoses.

Among the previously described species of Cis from southern Africa, only four of them were placed in species-groups: C. mooihoekite and C. pickeri (C. bilamellatus species-group); and C. chinensis and C. paraliacus (C. multidentatus species-group). The other previously described species were not placed in any group up to date, because it was necessary to describe the other new species at first and compare them to Cis from other biogeographic regions [10]. Here we place most described southern African species in species-groups. Cis muriceus and C. tessariplacus sp. n. are the only species that cannot be placed in any previously established group and we prefer not to propose new groups for them.

The following new species-groups are proposed: C. makrosoma species-group, C. neserorum species-group, C. regius species-group and C. westerncapensis species-group. The makrosoma group is characterized by: (i) anterocephalic edge simple or weakly bidentate; (ii) dual elytral punctation consisting of megapunctures forming more or less regular longitudinal rows, in-between rows filled with micropunctures bearing short bristles; (iii) very elongate, parallel-sided and flattened body; (iv) prosternum moderately long and flat. The makrosoma group includes the following species: the southern African C. makrosoma sp. n.; C. interpunctatus Mellié, 1849 from La Reunión, based on examination of material compared with type; C. cavifrons Blair, 1940 from Australia, doubtfully included based on the shape of male genitalia [3]. There are other Ethiopian species that may belong to this group, such as C. mahensis Scott, 1926 and C. parallelus Scott, 1926, but we did not have access to type of material or specimens compared with type. Both C. mahensis and C. parallelus can be synonyms of C. interpunctatus [26].

The neserorum group is characterized by: (i) anterocephalic edge and anterior pronotal edge barely to strongly emarginate, forming two short prominences or subtriangular plates in males; (ii) outer apical angle of male protibia projected in a tooth; (iii) dual and non-seriate (confuse) elytral punctation; (iv) pronotal and elytral vestiture indistinctly to distinctly dual. The neserorum group includes the following species: the southern African C. afer, C. aster, C. bicaesariatus sp. n., C. bimucronatus (doubtfully included), C. caffer, C. delagoensis, C. makebae, C. mandelai, C. masekelai, C. neserorum, C. stalsi and C. testaceus; and Cis renominatus Sandoval-Gómez, Lopes-Andrade & Lawrence, 2014 from Central Africa [27]. The eastern African species C. eichelbaumi Reitter, 1908 from Tanzania, C. pseudosphindus Reitter, 1908 from Tanzania and Kenya, and C. usambarinus Reitter, 1908 from Tanzania may belong to this group, but we not have access to the type or identified material of these species.

The regius group is characterized by: (i) head with a peculiar occipital tubercle close to vertex in males; (ii) pronotum with dual punctation, lateral to anterior edges broadly rounded and bearing a row of sparse setae; (iii) elytra with single punctation and vestiture of seriate setae. The unique southern African species included is C. regius, but the group also included C. biacutus Reitter, 1908 from Tanzania, Madagascar and Seychelles.

The westerncapensis group is characterized by: (i) anterocephalic edge and anterior pronotal edge simple; (ii) dual pronotal punctation; (iii) prosternum more or less carinate; (iv) elytral punctation dual and more or less seriate; (v) elytral vestiture single, subseriate to seriate, consisting of moderately short bristles arising from megapunctures. Included species: the southern African C. lacinipennis sp. n. and C. westerncapensis sp. n.

Most remaining southern African species are included in previously proposed groups, as follows: bilamellatus group (C. pickeri and C. mooihoekite) (for a definition of the group, see Lawrence [3] and Souza-Gonçalves & Lopes-Andrade [11]); comptus group (C. grobbelaarae sp. n.) (see Lawrence [6] and Kompantsev [28]); fuscipes group (C. capensis and C. fuscipes) (see Lawrence [3]); multidentatus group (C. chinensis and C. paraliacus) (see Lopes-Andrade [7] and Souza-Gonçalves & Lopes-Andrade [12]); and pacificus group (C. foveocephalus sp. n., C. mpumalangaensis sp. n., C. parvisetosus sp. n. and C. umlalaziensis sp. n.) (see Lawrence [3]).

3.1. Species Accounts

3.1.1. Cis afer Fåhraeus, 1871

Cis afer Fåhraeus 1871: 671 [21]. Type locality: Caffraria (=Kaffraria), currently Republic of South Africa: Eastern Cape Province (no specific locality); Ferrer 1997: 408 [24] {lectotype designation}.

Host fungi: Unknown.

Distribution: Ethiopian. Known from Eastern Cape Province (Republic of South Africa).

Comments: The species is known only from the type series [21]. There is no further record in the literature, as far as we have traced. This species is a member of the neserorum group.

3.1.2. Cis aster Souza-Gonçalves et Lopes-Andrade, 2017

Cis aster Souza-Gonçalves & Lopes-Andrade 2017: 340 [11]. Type locality: Republic of South Africa: KwaZulu-Natal Province, Twin Streams Nursery Forest.

Diagnosis: The species belongs to the neserorum group. It differs from other southern African species in the neserorum group (except for C. afer, C. bicaesariatus sp. n., C. caffer, C. makebae and C. masekelai) in males being devoid of concave impression in anterior pronotal portion. Cis aster differs from C. afer in males with comparatively longer pronotal plates; from C. bicaesariatus sp. n. in a comparatively longer body and larger prosternal process; and from C. caffer in males possessing much closer plates on anterocephalic and anterior pronotal edges. It differs from C. makebae and C. masekelai in males with first abdominal ventrite devoid of a sex patch.

Host fungi:Hexagonia tenuis (Hook.) Fr. (Polyporaceae), one record and possibly being breeding record [10].

Distribution. Ethiopian. Known from Twin Streams Nursery Forest (KwaZulu-Natal Province, Republic of South Africa).

Comments. The species is known only from the type series [11]. In the original description, it was mentioned that the species cooccured with the morphospecies Cis sp. Q. However, it was collected alone (corrigendum to Souza-Gonçalves and Lopes-Andrade [11]).

3.1.3. Cis bicaesariatus Souza-Gonçalves et Lopes-Andrade, sp. n.

ZooBank: http://www.zoobank.org/NomenclaturalActs/83B82A1D-EF13-4253-987F-6BAD2A972CCA

Figure 1(1–9); Figure 13

Cis sp. H in Neser [9].

Type locality: “Die Hel Nature Reserve”, coordinates 25°31’ S 29°48’ E (near Loskop Dam, Mpumalanga Province).

Etymology: The species name derives from the Latin noun “bi”, which means “two”, and “caesariatus”, which means “covered by hairs”, “long-haired”, both in the genitive singular. The name is a reference to the dual dorsal vestiture of the species.

Diagnosis: The species belongs to the neserorum group. It differs from other southern African species of the neserorum group (except for C. afer, C. aster, C. caffer, C. makebae and C. masekelai) in males being devoid of concave impression in anterior pronotal portion. Cis bicaesariatus sp. n. differs from C. afer, C. aster and C. caffer in a comparatively shorter body and in males bearing anterocephalic plates curved upwards and with rounded apex. It differs from C. makebae and C. masekelai in males with first abdominal ventrite devoid of a sex patch.

Description, male holotype (Figure 1(A–D)): Adult apparently not fully pigmented but in good condition, except for lacking three tarsi and for being covered in dust or fungus. Measurements in mm: TL 1.86, PL 0.66, PW 0.81, EL 1.20, EW 0.96, GD 0.71. Ratios: PL/PW 0.82, EL/EW 1.26, EL/PL 1.82, GD/EW 0.74, TL/EW 1.94. Body elongate, convex, dorsum reddish dark brown (except for dust-covered areas, but visible elsewhere in paratypes); venter reddish dark brown (visible only at abdominal ventrites due to dust-covering, but visible elsewhere in paratypes); antennae yellowish brown with club dark brown, palpi and tarsi yellowish brown; dorsal vestiture distinctly dual, consisting of suberect bristles from distinct lengths, easily discernible in high magnifications (>50×); ventral vestiture consisting of decumbent setae easily discernible in high magnifications (>50×). Head with anteriormost portion visible from above; dorsum with coarse and deep punctures, separated from each other by less than one puncture-width, with short decumbent bristle (0.01–0.02 mm) in each puncture; interspaces, microreticulate; anterocephalic edge produced and elevated forming two subrounded plates. Antennae with 10 antennomeres, lengths as follows (in mm, left antennae measured): 0.05, 0.04, 0.03, 0.02, 0.02, 0.02, 0.02, 0.06, 0.07, 0.10 (FL 0.11 mm, CL 0.22 mm, CL/FL 2.05). Eyes coarsely facetted, with about 90 ommatidia; GW 0.18 mm. Gula 0.55× as wide as head. Pronotum (Figure 1(D)) with coarse, deep, single punctation, devoid of impunctate median line; punctures separated from each other by one puncture-width or less; interspaces, microreticulate; vestiture distinctly dual, consisting of moderately long (~0.05 mm) and short (0.02–0.03 mm) suberect yellowish bristles; anterior edge produced and elevated forming two subtriangular plates; lateral edges crenulate, not explanate and not visible when seen from above; anterior corners barely angulate. Scutellar shield triangular, bearing few punctures and apparently glabrous; BW 0.11 mm; SL 0.09 mm. Elytra with non-seriate, dual punctation; megapunctures coarse, deep, about 3× as large as micropunctures, separated from each other by one megapuncture-width or less; interspaces a bit rugose; vestiture distinctly dual, consisting of moderately long (0.05–0.06 mm) and short (0.02–0.03 mm) suberect yellowish bristles, both arising from micropunctures. Metathoracic wings developed, apparently functional. Hypomera with fine, shallow punctation; each puncture bearing a fine decumbent seta; interspaces, microreticulate. Prosternum in front of coxae biconcave; interspaces, microreticulate. Prosternal process parallel-sided, relatively narrow, as long as prosternum at midline, apex truncate. Protibiae with maximum width about one-third of its length; apical edge devoid of spines; outer apical angle projected in acute tooth. Meso- and metatibiae without spines in apical edge. Metaventrite with fine, shallow punctures; interspaces, microreticulate; discrimen about one-third the length of metaventrite at midline. Abdominal ventrites with fine, shallow punctures, separated from each other by one puncture-width or less and bearing a fine yellowish decumbent seta; interspaces, microreticulate; length of ventrites (in mm, from base to apex at the longitudinal midline) as follows: 0,26, 0.12, 0.10, 0.10, 0.11; first abdominal ventrite devoid of sex patch. Male terminalia in a paratype (Figure 1(F–I)) with sternite VIII (Figure 1(F)) with posterior margin rounded, bearing short setae at middle and long at corners; anterio×r portion membranous. Tegmen (Figure 1(H)) 9.3× as long as wide; widest near apex; subparallel-sided; apex bilobed; apical portion membranous and rounded (Figure 1(H)), black arrows); anterior portion subtriangular. Basal piece (Figure 1(G)) triangular, as wide as long. Penis (Figure 1(I)) 1.4× as long as tegmen, 8.9× as long as wide; subparallel-sided; three acute angulations at apex (Figure 1(I), red arrows); shortly rounded emargination at anterior portion.

Females (Figure 1(E)): Anterior edge of head barely emarginate and anterior edge of pronotum rounded. Otherwise like males, but without pronotal and head plates and protibial tooth.

Variation: Females, measurements in mm (n = 4): TL 1.75–2.00 (1.87 ± 0.12), PL 0.53–0.65 (0.60 ± 0.06), PW 0.73–0.85 (0.79 ± 0.07), EL 1.20–1.35 (1.27 ± 0.07), EW 0.85–0.98 (0.91 ± 0.06), GD 0.65–0.78 (0.72 ± 0.06). Ratios: PL/PW 0.70–0.79 (0.76 ± 0.04), EL/EW 1.37–1.41 (1.39 ± 0.02), EL/PL 2.00–2.33 (2.12 ± 0.14), GD/EW 0.76–0.80 (0.79 ± 0.02), TL/EW 2.00–2.09 (2.05 ± 0.04).

Type material: Holotype: ♂ (SANC, dissected) “SOUTH AFRICA: MPU, Die Hel Nature Res., nr. Loskop Dam, 25°31’ S 29°48’ E, 10.viii.2008, S. & O.C. Neser\Ex bracket fungus Trametes sp., BF# 147\NATIONAL COLL.OF INSECTS Pretoria, South Africa\Cis bicaesariatus Souza-Gonçalves & Lopes-Andrade HOLOTYPUS [red paper]”. Paratypes: 5 ♀♀ as follows: 3 ♀♀ (2 CELC; 1 SANC) same data as the holotype; 1 ♀ (SANC) “SOUTH AFRICA: LIMP, Otter’ S Den 16 Km from Hoedspruit, 24°24’ S 30°49’ E, 18.vii.2008, D. van Heerden\Ex bracket fungus Coriolus versicolor, BF# 136\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♀ (SANC) “SOUTH AFRICA: NW, Castle Gorge, Magaliesberg, 25°49’ S 27°35’ E, 21.iv.2002, O.C. Neser\Adults emerged from bracket fungus Phaeolus schweinitzii, BF# 13\NATIONAL COLL. OF INSECTS Pretoria, South Africa”. All paratypes additionally labeled “Cis bicaesariatus Souza-Gonçalves & Lopes-Andrade PARATYPUS [yellow paper]”.

Host fungi:Phaeolus schweinitzii (Fr.) Pat. (Fomitopsidaceae), one record; Trametes sp., one record; Trametes versicolor (L.) Lloyd (Polyporaceae), one record.

Distribution: Ethiopian, known from northern Mpumalanga, southeastern Limpopo and northeastern North West (Republic of South Africa) (Figure 13).

Comments: This species was collected together with C. mandelai, C. mpumalangaensis sp. n., the invasive species Ceracis tabellifer (Mellié, 1849) and the parasitoid Astichus micans Neser 2012 (Hymenoptera: Eulophidae: Entiinae).

3.1.4. Cis bimucronatus Motschoulsky, 1851

Cis bimucronatus Motschoulsky 1851: 655 [25]. Type locality: Port Natal, currently Republic of South Africa: KwaZulu-Natal Province, Durban.

Host fungi: Unknown.

Distribution: Ethiopian. Known from Durban (KwaZulu-Natal Province, Republic of South Africa).

Comments: The species is known only from the type series [25]. There is no further record in the literature, as far as we have traced. The description of this species is anecdotal and the type has probably been lost. It is doubtfully included in the neserorum group.

3.1.5. Cis caffer Fåhraeus, 1871

Cis caffer Fåhraeus 1851: 670 [21]. Type locality: Caffraria (=Kaffraria), currently Republic of South Africa: Eastern Cape Province (no specific locality); Ferrer 1997: 408 [24] {lectotype designation}.

Host fungi: Unknown.

Distribution: Ethiopian. Known from Eastern Cape Province (Republic of South Africa).

Comments: The species is known only from the type series [21]. There is no further record in the literature, as far as we have traced. This species is a member of the neserorum group.

3.1.6. Cis capensis Mellié, 1849

Cis capensis Mellié 1849: 254 [22]. Type locality: Republic of South Africa: Western Cape Province, Cape of Good Hope.

Host fungi: Unknown.

Distribution: Ethiopian. Known from Cape of Good Hope (Western Cape Province, Republic of South Africa).

Comments: The species is known only from the type series [22]. There is no further record in the literature, as far as we have traced. This species is a member of the fuscipes group and may be even a synonym of C. fuscipes properly, a hypothesis that shall be evaluated after careful examination of the type of C. capensis.

3.1.7. Cis chinensis Lawrence, 1991

Cis chinensis Lawrence 1991: 288 [29]. Type locality: China (no specific locality); Madenjian et al. 1993: 47 [30] {found in fungi imported from China to USA, not free-living}; Jinachai et al. 2002 [31] {record from Thailand}; Buder et al. 2008: 171 [4] {GenBank access numbers: FM877940, FM877793, FM877874}; Jelínek 2008: 56 [32] {listed among Palearctic species}; Lopes-Andrade 2008: 36 [7] {record from Brazil}; Rose 2009: 282 [33] {record from France and La Réunion}; Reibnitz & Kunz 2011: 45 [34] {record from Baden-Württemberg, Germany, and Hungary}; Reibnitz 2012 [35] {distribution}; Rose 2012: 346 [36] {host-fungi and record from Launaguet, France}; Diéguez Fernandéz 2013: 104 [37] {record from Spain}; Rose 2014: 1 [38] {record from France}; Amini et al. 2016 [39] {record from Iran}; Lawrence 2016: 42 [3] {redescription and record from Australia}; Rose & Zagatti 2016: 292, 302, 304–305 [40] {host-fungi in France}; Németh et al. 2017: 28 [41] {record from Budapest, Hungary}; Souza-Gonçalves & Lopes-Andrade 2018: 503 [13] {record from Republic of South Africa and Northern Mariana Islands}.

Cis multidentatus (Pic 1917) sensu Lohse & Reibnitz 1991: 104 [42] {reported from Italy and Germany}; Jelínek 2008: 57 [32] {records from Italy and Germany}; Lopes-Andrade 2008: 42 [7] {reported from Malta}; Shugran et al. 2018: 42 [43] {reported from Iraq}.

Plesiocis sp. sensu Yan et al. 1998 [44] {record from Shandong province, China}.

Diagnosis: The species belongs to the multidentatus group. It differs from C. paraliacus in bearing elytral punctation single and lateral pronotal edges barely to completely visible from above.

Host fungi:Agaricus sp. (Agaricaceae), one record [42]; Coprinus sp. (Agaricaceae), one record [42]; Daedaleopsis nitida (Durieu et Mout.) Zmitr. et Malysheva (Polyporaceae), one record [37]; Fomitopsis pinicola (Sw.) P. Karst. (Fomitopsidaceae), one record [37]; Ganoderma lucidum (Ganodermataceae), seven records as pest of commercial dried fungi [3,29,30,31,32,42]; Gloeophyllum abietinum (Bull.) P. Karst. (Gloeophyllaceae), one record [37]; Lactarius sp. (Russulaceae), one record [41]; Pleurotus ostreatus (Jacq.) P. Kumm. (Pleurotaceae), one record [41]; Russula sp. (Russulaceae), one record [41]; Schizophyllum commune Fr. (Schizophyllaceae), one breeding record [7]; Trametes sp. (Polyporaceae), one breeding record [42]; Trametes gibbosa (Pers.) Fr., one record [40].

Distribution: Cosmopolitan. Known from east and southeast Asia, northeastern USA, southeastern Brazil, southern, western and eastern Europe, Caucasus, Western Indian Ocean and southern Africa (southern Western Cape, Republic of South Africa). Recorded as pest of the fungus Ganoderma lucidum (known as Reishi in Japan and Ling Zhi in China) in east and Southeast Asia [29,30,31] and Australia [3].

Comments: This species was collected together with C. pickeri and the invasive species Cer. tabellifer [13].

3.1.8. Cis delagoensis Pic, 1916

Cis delagoensis Pic 1916: 14 [23]. Type locality: Mozambique: Delagoa Bay (=Maputo Bay).

Host fungi: Unknown.

Distribution: Ethiopian. Known from Maputo Bay (Mozambique).

Comments: The species is known only from the type series [23]. There is no further record in the literature, as far as we have traced. There are identified specimens in the Natural History Museum (NHM) (pictures examined by us) from Democratic Republic of the Congo, Sierra Leone and Uganda, but we are not sure they are conspecific. This species is a member of the neserorum group.

3.1.9. Cis foveocephalus Souza-Gonçalves et Lopes-Andrade, sp. n.

ZooBank: http://www.zoobank.org/NomenclaturalActs/DD6E566F-C3A5-40BE-8528-582903A57E26

Figure 2(A–I); Figure 14.

Cis sp. P in Neser [9].

Type locality: “Mooihoek Farm” (near Wakkerstroom), coordinates 27°13’ S 30°32’ E (Pixley Ka Seme Local Municipality, Gert Sibande District, Mpumalanga Province).

Etymology: The species name is derived from the Latin noun “foveolae”, which means “small pit”, and the latinized Greek noun “kephale”, which means “head”, both in the genitive singular. The name is a reference to the sex patch present on the head of males.

Diagnosis: The species belongs to the pacificus group. It differs from all other southern African species of the pacificus group in males bearing a sex patch in vertex as well as in first abdominal ventrite.

Description, male holotype (Figure 2(A–D)): Adult fully pigmented and in good conditions, except for lacking one leg, five tarsi and both antennae. Measurements in mm: TL 1.75, PL 0.60, PW 0.76, EL 1.15, EW 0.83, GD 0.62. Ratios: PL/PW 0.78, EL/EW 1.39, EL/PL 1.93, GD/EW 0.75, TL/EW 2.11. Body elongate, convex, dorsum and venter reddish dark brown; palpi and tarsi yellowish brown; dorsal vestiture single, consisting of short suberect bristles, easily discernible in high magnifications (>50×); ventral vestiture of decumbent setae easily discernible in high magnifications (>50×). Head with anteriormost portion visible from above; dorsum with coarse and deep punctures, separated from each other by one puncture-width or less, with short decumbent bristle (0.01–0.02 mm) arising from each puncture; interspaces, microreticulate; vertex bearing setose sex patch with transverse diameter of 0.06 mm; anterocephalic edge bearing two small triangular tubercles. Antennae with ten antennomeres, lengths as follows (in mm, left antennae measured in a paratype): 0.05, 0.04, 0.05, 0.04, 0.03, 0.03, 0.02, 0.05, 0.05, 0.06 (FL 0.15 mm, CL 0.15 mm, CL/FL 1.00). Eyes coarsely facetted, with about 60 ommatidia; GW 0.15 mm. Gula 0.43× as wide as head. Pronotum (Figure 2(D)) with irregularly distributed, dual punctation, devoid of impunctate median line; megapunctures coarse, deep, about 2× as large as micropunctures, separated from each other by two megapuncture-widths or less; interspaces, microreticulate; vestiture single, consisting of short suberect yellowish bristles (0.01–0.02 mm) arising from megapunctures; anterior edge rounded; lateral edges crenulate, not explanate and not visible when seen from above; anterior corners rounded. Scutellar shield triangular, bearing few punctures and few bristles; BW 0.07 mm; SL 0.06 mm. Elytra with non-seriate, dual punctation; megapunctures coarse, deep, about 2× as large as micropunctures, separated from each other by two megapuncture-widths or less; interspaces a bit rugose; vestiture single, consisting of short suberect yellowish bristles (0.01–0.02 mm) arising from megapunctures. Metathoracic wings developed, apparently functional. Hypomera with coarse, shallow punctation; each puncture bearing a fine decumbent seta; interspaces, microreticulate. Prosternum in front of coxae biconcave and barely carinate; interspaces, microreticulate. Prosternal process subparallel-sided, about 0.9× as long as prosternum at midline, apex rounded. Protibiae with maximum width about one-third of its length; apical edge devoid of spines; outer apical angle projected in acute tooth. Meso- and metatibiae without spines in apical edge. Metaventrite with coarse, deep punctures; interspaces, microreticulate; discrimen about two-fifths the length of metaventrite at midline. Abdominal ventrites with coarse, moderately deep punctures, separated from each other by one puncture-width or less and bearing a fine decumbent yellowish seta; interspaces, microreticulate; length of ventrites (in mm, from base to apex at the longitudinal midline) as follows: 0.27, 0.11, 0.09, 0.08, 0.10; first abdominal ventrite bearing margined, circular, setose sex patch posterad of center, with transverse diameter of 0.05 mm. Male terminalia in a paratype (Figure 2(F–I)) with sternite VIII (Figure 2(F)) with posterior margin emarginate, bearing short setae at middle and long setae at acute corners; anterior portion membranous. Tegmen (Figure 2(H)) 3.8× as long as wide, widest near apex; subparallel-sided but more or less sinuate; one rounded emargination in each side forming three lobes at apex, the lateral ones short and acute (Figure 2(H), big black arrows) and the mid one long, somewhat arrow-shaped and covered by sensillae (Figure 2(I), small black arrow); anterior portion triangular. Basal piece (Figure 2(G)) oval, 1.3× as long as wide. Penis (Figure 2(I)) 0.9× as long as tegmen, 5.3× as long as wide; subparallel-sided and bearing sensillae near to membranous apex; anterior portion rounded.

Females (Figure 2(E)): Anterior edge of head truncate and anterior edge of pronotum rounded. Otherwise like males, but devoid of cephalic tubercles, abdominal and cephalic sex patch, and protibial tooth.

Variation: Males, measurements in mm (n = 9, including the holotype): TL 1.43–1.83 (1.65 ± 0.13), PL 0.48–0.65 (0.56 ± 0.06), PW 0.58–0.78 (0.67 ± 0.08), EL 0.95–1.23 (1.09 ± 0.08), EW 0.65–0.88 (0.76 ± 0.07), GD 0.48–0.62 (0.54 ± 0.05). Ratios: PL/PW 0.77–0.96 (0.83 ± 0.07), EL/EW 1.31–1.53 (1.44 ± 0.07), EL/PL 1.77–2.15 (1.58 ± 0.13), GD/EW 0.63–0.77 (0.71 ± 0.04), TL/EW 2.06–2.29 (2.17 ± 0.08). Females, measurements in mm (n = 6): TL 1.30–1.78 (1.61 ± 0.10), PL 0.50–0.55 (0.52 ± 0.02), PW 0.60–0.68 (0.64 ± 0.03), EL 1.00–1.23 (1.10 ± 0.08), EW 0.70–0.78 (0.76 ± 0.03), GD 0.58–0.63 (0.58 ± 0.02). Ratios: PL/PW 0.74–0.88 (0.81 ± 0.05), EL/EW 1.33–1.58 (1.45 ± 0.09), EL/PL 2.00–2.23 (2.12 ± 0.10), GD/EW 0.74–0.82 (0.77 ± 0.04), TL/EW 2.00–2.29 (2.13 ± 0.13).

Type material: Holotype: ♂ (SANC) “SOUTH AFRICA: MPU, Mooihoek Farm, nr. Wakkerstroom, 27°13’ S 30°32’ E, 15.vii.2008. O & S Neser\Ex bracket fungus Stereum ostrea, BF# 138\NATIONAL COLL. OF INSECTS Pretoria, South Africa\Cis foveocephalus Souza-Gonçalves & Lopes-Andrade HOLOTYPUS [red paper]”. Paratypes: 11 ♂♂ and 11 ♀♀ as follows: 6 ♂♂ (2 CELC, dissected; 4 SANC) and 4 ♀♀ (2 CELC; 2 SANC) same data as the holotype; 2 ♂♂ (1 CELC; 1 SANC) and 6 ♀♀ (2 CELC; 4 SANC) “SOUTH AFRICA: MPU, Mooihoek Farm, nr. Wakkerstroom, 27°13’ S 30°32’ E, 15.vii.2008, O & S Neser\Ex bracket fungus, BF# 107\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♂ (CELC) “SOUTH AFRICA: NW, Groblerskloof, nr. Buffelspoort Dam, 25°51’ S 27°26’ E, 07.viii.2008, S. Neser\Ex bracket fungus Trametes cingulata, BF# 146”; 1 ♂ (SANC) “SOUTH AFRICA: NATAL, Cathedral Peak Forestry Area, 28°55’ S 29°14’ E, 10.xi.1981, SJv Tonder & C Kok \ NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♂ and 1 ♀ (CELC) “SOUTH AFRICA: NW, Cave Kloof, Magaliesberg, nr. Bufflspoort Dam, 25°51’ S 27°26’ E, 24.vii.2008, S. Neser\Ex bracket fungus, BF# 103\Ex bracket fungus Ganoderma applanatum, BF# 103\NATIONAL COLL. OF INSECTS Pretoria, South Africa”. All paratypes additionally labeled “Cis foveocephalus Souza-Gonçalves & Lopes-Andrade PARATYPUS [yellow paper]”.

Host fungi:Ganoderma applanatum (Pers.) Pat., one record; Stereum ostrea (Blume & T. Nees) Fr., one breeding record; Tramates cingulata Berk., one record.

Distribution: Ethiopian. Known from southeastern Mpumalanga, northeastern North West and western KwaZulu-Natal (Republic of South Africa) (Figure 14).

Comments: This species was collected together with C. mpumalangaensis sp. n., C. tessariplacus sp. n., the invasive species Cer. tabellifer and the parasitoid A. naiadis Neser, 2012.

3.1.10. Cis fuscipes Mellié, 1849

Figure 3(A–D); Figure 15.

Cis fuscipes Mellié 1849: 271, pl. 10, Figure 4(A) [22]. Type locality: USA: Massachusetts, Boston; Lawrence 1967: 1–14 [45] {synonyms, distribution and biology}; Lawrence 1971: 460 [6] {distribution and host fungi at North America}; Jelínek 2008: 57 [32] {listed among Paleartic species}; Lawrence 2016: 53 [3] {redescription and record from Australia}; Lopes-Andrade et al. 2016: 347 [46] {record from New Brunswick, Canada}.

Cis atripennis Mellié 1849: 258 [22]; Lawrence 1967: 11 [45] {synonym}.

Cis chevrolatii Mellié 1849: 249 [22]; Lawrence 1967: 11 [45] {synonym}.

Cis dubius Mellié 1849: 273 [22]; Lawrence 1967: 11 [45] {synonym}.

Cis carolinae Casey 1898: 78 [47]; Lawrence 1967: 11 [45] {synonym}.

Cis impressa Casey 1898: 79 [47]; Lawrence 1967: 11 [45] {synonym}.

Cis pallens Casey 1898: 78 [47]; Lawrence 1967: 11 [45] {synonym}.

Diagnosis: The species belongs to the fuscipes group. It differs from C. capensis in males bearing truncate or weakly convex anterocephalic edge and anterior pronotal edge rounded, but only female C. fuscipes have been recorded from southern Africa. It differs from females of C. capensis in the anteriormost portion of head visible when see from above.

Additional material: 6 ♀♀ as follows: 5 ♀♀ (2 CELC, dissected; 3 SANC, one dissected) “SOUTH AFRICA WCape, Garden of Eden Indig. Forest, nr. Knysna, 34°02’ S 23°12’ E, 1.iii.1991, AJ Hendricks\Emerged from log on forest floor of Olea capensis macrocarpa OLEACEAE, UA679\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♀ (SANC) “SOUTH AFRICA: MPU, Mooihoek Farm, nr. Wakkerstroom, 27°13’ S 30°32’ E, 15.vii.2008, O & S Neser\Ex bracket fungus, BF# 129\Ex bracket fungus Coriolus versicolor, BF# 129\NATIONAL COLL. OF INSECTS Pretoria, South Africa”. All additionally labeled “Cis fuscipes Mellié, 1849; I. Souza-Gonçalves & C. Lopes-Andrade det.”.

Host fungi:Bjerkandera adusta (Willd.) P. Kasrt. (Merulliaceae), one breeding record [6]; Cerioporus squamosus (Huds.) Quél. (Polyporaceae), one breeding record [6,45]; Daedalea ambigua Berk. (Fomitopsidaceae), one record [6]; Fomitopsis pinicola (Sw.) P. Karst., one record [6]; Ganoderma applanatum (Pers.) Pat., one record [6]; Ganoderma brownii (Murrill) Gilb. (Ganodermataceae), one breeding record [6,45]; Lenzites betulinus (L.) Fr. (Polyporaceae), 12 records, three being breeding records [6]; Perenniporia fraxinophila (Peck) Ryvarden, one record [6,45]; Phellinus gilvus (Schwein.) Pat. (Hymenochaetaceae), one record [45]; Poronidulus conchifer (Schwein.) Murrill (Polyporaceae), three records, two being breeding records [6,45]; Trametes hirsuta (Wulfen) Lloyd, 14 records, 10 being breeding records [6,45]; Trametes pubescens (Schumach.) Pilát (Polyporaceae), five records, one being breeding record [6]; Trametes suaveolens (L.) Fr. (Polyporaceae), one record [45]; Trametes subectypa (Murrill) Gilb. & Ryvarden (Polyporaceae), one record [6]; Trametes versicolor (L.) Lloyd, 93 records, 48 being breeding records [3,6,45].

Distribution: Cosmopolitan. Known from the Holarctic kingdom, widely distributed in North America. Females have been previpously recorded from Australia, Cuba, Hawaii, Madeira and New Zealand, where they were probably introduced [3]. The additional material was collected in southeastern Mpumalanga and southern Western Cape (Republic of South Africa) (Figure 15).

Comments: This species was collected together with C. mooihoekite, C. neserorum, C. parvisetosus sp. n., C. pickeri, the morphospecies Cis sp. S, Cis sp. Y and the invasive species Cer. tabellifer in South Africa. Only females of C. fuscipes were collected by the staff of SANC (Figure 3(A–D)), suggesting that this species may be represented only by parthenogenetic populations in Republic of South Africa.

3.1.11. Cis grobbelaarae Souza-Gonçalves et Lopes-Andrade, sp. n.

ZooBank: http://www.zoobank.org/NomenclaturalActs/FFEC3023-8443-4DAC-B983-29E0F9C32486

Figure 4(A–I); Figure 14.

Cis sp. C in Neser [9].

Type locality: “Die Hel Nature Reserve”, coordinates 25°31’ S 29°48’ E (near Loskop Dam, Mpumalanga Province).

Etymology: The new species is named in honor of the South African taxonomist Elizabeth Grobbelaar, who collected all paratypes from D’Nyala Nature Reserve. The species name is Latinized from “Grobbelaar” using the feminine suffix in the genitive singular (-ae).

Diagnosis: The species belongs to the comptus group. It differs from all other southern African species (except for C. makrosoma sp. n.) by the pattern of the dual elytral punctation, consisting of megapunctures forming more or less regular longitudinal rows, in-between rows filled with micropunctures bearing short bristles. It differs from C. makrosoma sp. n. in males with anterocephalic edge bearing small angulations, less elongated and more convex body, and a comparatively shorter and slightly tumid prosternum.

Description, male holotype (Figure 4(A–D)): Adult fully pigmented and in good conditions. Measurements in mm: TL 1.78, PL 0.57, PW 0.71, EL 1.20, EW 0.82, GD 0.62. Ratios: PL/PW 0.80, EL/EW 1.46, EL/PL 2.10, GD/EW 0.75, TL/EW 2.16. Body elongate, convex, dorsum and venter reddish dark brown; antennae, palpi and tarsi yellowish brown; dorsal vestiture single, consisting of moderately short suberect bristles, easily discernible in low magnifications (<50×); ventral vestiture of decumbent setae easily discernible in low magnifications (<50×). Head with anteriormost portion visible from above; dorsum with coarse and deep punctures, separated from each other by one puncture-width or less, with short decumbent bristle (0.01–0.02 mm) arising from each puncture; interspaces, microreticulate; anterocephalic edge with four small angulations (barely discernible). Antennae with 10 antennomeres, lengths as follows (in mm, left antennae measured): 0.07, 0.04, 0.04, 0.03, 0.02, 0.02, 0.02, 0.04, 0.04, 0.06 (FL 0.13 mm, CL 0.15 mm, CL/FL 1.10). Eyes coarsely facetted, with about 60 ommatidia; GW 0.16 mm. Gula 0.38× as wide as head. Pronotum (Figure 4(D)) with coarse, deep, single punctation, devoid of impunctate median line; punctures distributed irregularly, separated from each other by one puncture-width or less; interspaces, microreticulate; vestiture single, consisting of moderately short suberect yellowish bristles (0.03–0.04 mm); anterior edge rounded; lateral edges crenulate, explanate and completely visible when seen from above; anterior corners rounded. Scutellar shield triangular, bearing few punctures and apparently glabrous; BW 0.14 mm; SL 0.08 mm. Elytra with seriate, dual punctation; megapunctures coarse, deep, about 2× as large as micropunctures, separated from each other by one megapuncture-width or less, forming more or less longitudinal rows, in-between rows filled with micropunctures; interspaces a bit rugose; vestiture single, consisting of moderately short suberect yellowish bristles (0.03–0.04 mm) arising from micropunctures. Metathoracic wings developed apparently functional. Hypomera with coarse, shallow punctation; each puncture bearing a fine decumbent seta; interspaces, microreticulate. Prosternum in front of coxae slightly tumid; interspaces, microreticulate. Prosternal process subparallel-sided, as long as prosternum at midline, apex rounded. Protibiae with maximum width about one-fourth of its length; apical edge devoid of spines; outer apical angle projected in acute tooth. Meso- and metatibiae without spines in apical edge. Metaventrite with coarse, deep punctures; interspaces, microreticulate; discrimen about 0.5× the length of metaventrite at midline. Abdominal ventrites with coarse, moderately deep punctures, separated from each other by one puncture-width or less and bearing a fine decumbent seta; interspaces, microreticulate; length of ventrites (in mm, from base to apex at the longitudinal midline) as follows: 0.24, 0.10, 0.09, 0.09, 0.12; first abdominal ventrite bearing unmargined, small, almost glabrous, sex patch anterad of center, with transverse diameter of 0.04 mm. Male terminalia in a paratype (Figure 4(F–I)) with sternite VIII (Figure 4(F)) with the posterior margin barely emarginate, bearing short setae at middle and long setae at rounded corners; anterior portion membranous. Tegmen (Figure 4(H)) 1.9× as long as wide, widest at apex; sides expanding from basal third to apex; apex with one deep emargination in each side and with acute angulations at corners (Figure 4(H), black arrows); anterior portion rounded. Basal piece (Figure 4(G)) semicircular, 1.6× as wide as long. Penis (Figure 4(I)) 0.6× as long as tegmen, 3× as long as wide; sides expanding until basal two-thirds and then converging to apex; apical portion membranous and with sclerotization at middle, apex truncate (Figure 4(I), red arrows); anterior portion with broadly rounded, deep emargination.

Females (Figure 4(E)): Anterior edge of head truncate and edge of pronotum rounded. Otherwise like males, but devoid of abdominal sex patch and protibial tooth.

Variation: Males, measurements in mm (n = 8, including the holotype): TL 1.38–1.93 (1.73 ± 0.17), PL 0.43–0.63 (0.56 ± 0.06), PW 0.60–0.80 (0.70 ± 0.06), EL 0.95–1.33 (1.18 ± 0.11), EW 0.68–0.88 (0.80 ± 0.07), GD 0.53–0.70 (0.59 ± 0.06). Ratios: PL/PW 0.71–0.86 (0.80 ± 0.06), EL/EW 1.41–1.65 (1.48 ± 0.08), EL/PL 2.00–2.29 (2.12 ± 0.10), GD/EW 0.69–0.90 (0.75 ± 0.07), TL/EW 2.03–2.45 (2.18 ± 0.14). In small males, the angulations at the anterocephalic edge are barely distinguishable. In large males, there are four angulations at the anterocephalic edge (two at corners and two at middle). Females, measurements in mm (n = 7): TL 1.68–2.08 (1.88 ± 0.14), PL 0.53–0.68 (0.60 ± 0.06), PW 0.65–0.85 (0.76 ± 0.07), EL 1.15–1.40 (1.28 ± 0.09), EW 0.75–0.95 (0.88 ± 0.28), GD 0.58–0.75 (0.67 ± 0.06). Ratios: PL/PW 0.74–0.81 (0.79 ± 0.03), EL/EW 1.39–1.53 (1.46 ± 0.06), EL/PL 1.92–2.33 (2.14 ± 0.07), GD/EW 0.72–0.79 (0.76 ± 0.02), TL/EW 2.03–2.23 (2.14 ± 0.07).

Type material: Holotype: ♂ (SANC) “SOUTH AFRICA: MPU, Die Hel Nature Res., nr. Loskop Dam, 25°31’ S 29°48’ E, 10.viii.2008, S. & O.C. Neser\Ex unidentified bracket fungus, BF# 122\NATIONAL COLL. OF INSECTS Pretoria, South Africa\Cis grobbelaarae Souza-Gonçalves & Lopes-Andrade HOLOTYPUS [red paper]”. Paratypes: 22 ♂♂ and 14 ♀♀ as follows: 4 ♂♂ (1 CELC, dissected; 3 SANC) and 4 ♀♀ (1 CELC; 3 SANC) same data as the holotype; 1 ♂ and 1 ♀ (CELC) “SOUTH AFRICA: MPU, Die Hel Nature Res., nr. Loskop Dam, 25°31’ S 29°48’ E, 10.viii.2008, S. & O.C. Neser\Ex bracket fungus Trametes meyenii, BF# 163\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 7 ♂♂ (2 CELC, one dissected; 5 SANC) and 1 ♀ (SANC) “SOUTH AFRICA: MPU, Die Hel Nature Res., nr. Loskop Dam, 25°31’ S 29°48’ E, 10.viii.2008, S. & O.C. Neser\Ex bracket fungus, BF# 120\Ex bracket fungus Coriolopsis polyzona, BF# 120\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 6 ♂♂ (2 CELC; 4 SANC) and 5 ♀♀ (2 CELC; 3 SANC) “SOUTH AFRICA: MPU, Die Hel Nature Res., nr. Loskop Dam, 25°31’ S 29°48’ E, 10.viii.2008, S. & O.C. Neser\Ex bracket fungus, BF# 119\Ex bracket fungus Trametes sp., BF# 119\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 4 ♂♂ (2 CELC, one dissected; 2 SANC) and 3 ♀♀ (1 CELC; 2 SANC) “SOUTH AFRICA: TvL, D’Nyala Nature Res., near Ellisras, 23.45S 27.49E, 850 m, 29.ix.1989, E. Grobbelaar\Collected from bracket fungus body\NATIONAL COLL. OF INSECTS Pretoria, S. Afr.”. All paratypes additionally labeled “Cis grobbelaarae Souza-Gonçalves & Lopes-Andrade PARATYPUS [yellow paper]”.

Host fungi:Trametes sp. (Polyporaceae), one record; Trametes meyenii (Klotzsch) Lloyd (Polyporaceae), one record; Trametes polyzona (Pers.) Ryvarden (Polyporaceae), one record.

Distribution: Ethiopian. Known from northern Mpumalanga and northwestern Limpopo (Republic of South Africa) (Figure 14).

Comments: This species was collected together with C. neserorum, C. makebae, C. masekelai, C. mandelai, C. mooihoekite, the morphospecies Cis sp. Y, the invasive species Cer. tabellifer and the parasitoid A. micans.

3.1.12. Cis lacinipennis Souza-Gonçalves et Lopes-Andrade, sp. n.

ZooBank: http://www.zoobank.org/NomenclaturalActs/36C95C72-DF38-4BAB-A141-216679FA3D58

Figure 5(A–H); Figure 13.

Type locality: “Strathedene Farm” (near Nottingham Road), coordinates 29°21’ S 30°01’ E (uMngeni Local Municipality, uMgungundlovu District, KwaZulu-Natal Province).

Etymology: The species name derives from the Latin noun “lacina”, which means “flap”, and “penis”, both in the genitive singular. The name is a reference to the shape of the penis of this species, which bears a flap in each side of base.

Diagnosis: The species belongs to the westerncapensis group. It differs from C. westerncapensis sp. n. in bearing slightly shorter and seriate vestiture (0.02–0.03 mm), darker dorsal coloration and a comparatively narrower body.

Description, male holotype (Figure 5(A–D)): Adult fully pigmented and in good conditions, except for lacking one tarsus. Measurements in mm: TL 1.25, PL 0.40, PW 0.50, EL 0.85, EW 0.56, GD 0.48. Ratios: PL/PW 0.81, EL/EW 1.53, EL/PL 2.11, GD/EW 0.86, TL/EW 2.26. Body elongate, convex, dorsum and venter reddish dark brown; antennae yellowish brown with club dark brown, palpi and tarsi yellowish brown; dorsal vestiture single, consisting of moderately short suberect bristles, easily discernible in high magnifications (>75×); ventral vestiture of decumbent setae easily discernible in high magnifications (>75×). Head with anteriormost portion visible from above; dorsum with coarse and deep punctures, separated from each other by one puncture-width or less, with decumbent (~0.02 mm) minute bristle arising from each puncture; interspaces, microreticulate; anterocephalic edge truncate. Antennae with 10 antennomeres, lengths as follows (in mm, left antennae measured): 0.05, 0.04, 0.03, 0.02, 0.02, 0.02, 0.02, 0.03, 0.03, 0.04 (FL 0.10 mm, CL 0.11 mm, CL/FL 1.07). Eyes coarsely facetted, with about 50 ommatidia; GW 0.11 mm. Gula 0.46× as wide as head. Pronotum (Figure 5(D)) with irregularly distributed, dual punctation bearing an impunctate median line beginning around three punctures-width of base until disc; megapunctures coarse, deep, about 2× as large as micropunctures, separated from each other by one megapuncture-width or less; interspaces, microreticulate; vestiture single, consisting of moderately short suberect yellowish bristles (0.02–0.03 mm) arising from megapunctures; anterior edge rounded; lateral edges barely crenulate, not explanate and not visible when seen from above; anterior corners rounded. Scutellar shield triangular, bearing few punctures and few bristles; BW 0.07 mm; SL 0.05 mm. Elytra with subseriate, dual punctation; megapunctures coarse, deep, about 2× as large as micropunctures, separated from each other by two megapuncture-widths or less; interspaces a bit rugose; vestiture seriate, single, consisting of moderately short suberect yellowish bristles (0.02–0.03 mm) arising from megapunctures. Metathoracic wings developed, apparently functional. Hypomera with coarse, shallow punctation; each puncture bearing one fine decumbent seta; interspaces, microreticulate. Prosternum in front of coxae biconcave and barely carinate; interspaces, microreticulate. Prosternal process subparallel-sided, about 0.9× as long as prosternum at midline, apex rounded. Protibiae with maximum width about one-fourth of its length; apical edge devoid of spines; outer apical angle projected in acute tooth. Meso- and metatibiae without spines in apical edge. Metaventrite with coarse, deep punctures; interspaces, microreticulate; discrimen about one-third the length of metaventrite at midline. Abdominal ventrites with coarse, moderately deep punctures, separated from each other by one puncture-width or less and bearing one fine decumbent yellowish seta; interspaces, microreticulate; length of ventrites (in mm, from base to apex at the longitudinal midline) as follows: 0.18, 0.07, 0.05, 0.06, 0.08; first abdominal ventrite bearing unmargined, large, circular setose sex patch at middle, with transverse diameter of 0.07 mm. Male terminalia in a paratype (Figure 5(F–H)) with sternite VIII (Figure 5(F)) with posterior margin barely emarginate, bearing short setae at middle and long setae at subacute corners; anterior portion membranous. Tegmen (Figure 5(G)) 1.7× as long as wide, widest at apical third; sides expanding from basal third to apex; apex with rounded emargination and two small tubercles at middle (Figure 5(G), black arrows); anterior portion triangular. Penis (Figure 5(H)) as long as tegmen, 2.4× as long as wide; subparallel-sided; apex rounded; flaps (Figure 5(H), red arrows) and rounded emargination at anterior portion.

Females (Figure 5(36)): Anterior edge of head truncate and anterior edge of pronotum rounded. Otherwise like males, but devoid of abdominal sex patch and protibial tooth.

Variation: Males, measurements in mm (n = 12, including the holotype): TL 1.15–1.38 (1.29 ± 0.07), PL 0.38–0.45 (0.41 ± 0.02), PW 0.43–0.53 (0.48 ± 0.04), EL 0.78–0.95 (0.88 ± 0.09), EW 0.50–0.63 (0.58 ± 0.04), GD 0.40–0.53 (0.47 ± 0.04). Ratios: PL/PW 0.80–0.94 (0.85 ± 0.04), EL/EW 1.41–1.71 (1.54 ± 0.11), EL/PL 2.06–2.40 (2.18 ± 0.12), GD/EW 0.73–0.95 (0.82 ± 0.06), TL/EW 2.09–2.50 (2.25 ± 0.14). Females, measurements in mm (n = 11): TL 1.15–1.50 (1.32 ± 0.11), PL 0.38–0.50 (0.42 ± 0.04), PW 0.45–0.65 (0.51 ± 0.06), EL 0.78–1.00 (0.90 ± 0.07), EW 0.55–0.70 (0.61 ± 0.05), GD 0.45–0.58 (0.51 ± 0.04). Ratios: PL/PW 0.75–0.89 (0.83 ± 0.04), EL/EW 1.35–1.59 (1.47 ± 0.07), EL/PL 2.00–2.33 (2.14 ± 0.13), GD/EW 0.78–0.91 (0.84 ± 0.04), TL/EW 2.00–2.27 (2.16 ± 0.08).

Type material: Holotype: ♂ (SANC) “SOUTH AFRICA: KZN, Strathdene Farm, nr. Nottingham Rd., 29°21’ S 30°01’ E, 13.vii.2008, S & OC Neser\Ex bracket fungus, BF# 124\NATIONAL COLL. OF INSECTS Pretoria, South Africa\Cis lacinipennis Souza-Gonçalves & Lopes-Andrade HOLOTYPUS [red paper]”. Paratypes: 18 ♂♂ and 19 ♀♀ as follows: 8 ♂♂ (3 CELC, 2 dissected; 5 SANC) and 12 ♀♀ (4 CELC; 8 SANC) same data as the holotype; 1 ♂ and 1 ♀ (SANC) “Grootvadersbos, J.K. Grobler, 22.8.1956, Ac.X.846 \ NATIONAL COLL. OF INSECTS Pretoria, S. Afr.”; 5 ♂♂ (1 CELC, dissected; 4 ANIC) and 2 ♀♀ (1 CELC; 1 ANIC) “REP. SOUTH AFRICA: Natal, 75 Km WSW Estcourt Cathedral Peaks For. Sta., 7-31.XII.79, S. & J. Peck \ Ber 28, 29.XII.79, Podocarp forest rotted wood w/fungi, 1500 m”; 2 ♂♂ and 1 ♀ (ANIC) “REP. SOUTH AFRICA: Natal, 75 Km WSW Estcourt Cathedral Peaks For. Sta., 7-31.XII.79, S. & J. Peck\Ber 5, 14.XII.79, Podocarp forest rotted wood, bark, fungi, 1500 m”; 1 ♂ and 1 ♀ (CELC) “REP. SOUTH AFRICA: Natal, 75 Km WSW Estcourt Cathedral Peaks For. Sta., 7-31.XII.79, S. & J. Peck\Ber 6, 15.XII.79, Podocarp forest litter, mossy, boulder bases, 1500m”; 1 ♀ (1 ANIC) “REP. SOUTH AFRICA: Natal, 75 km WSW Estcourt Cathedral Peaks For. Sta., 7-31.XII.79, S. & J. Peck\Ber 1, 10.XII.79, sifted moss, Podocarp forest, 1500m”; 1 ♀ (ANIC) “REP. SOUTH AFRICA: Natal, 75 km WSW Estcourt Cathedral Peaks For. Sta., 7-31.XII.79, S. & J. Peck\Rainbow Gorge, Podocarp For., 1500 m, malaise traps 8, 13.XII.79”. All paratypes additionally labeled “Cis lacinipennis Souza-Gonçalves & Lopes-Andrade PARATYPUS [yellow paper]”.

Host fungi: Unknown.

Distribution: Ethiopian. Known from western KwaZulu-Natal and northern Free State (Republic of South Africa) (Figure 13).

Comments. This species was collected together with the invasive species Cer. tabellifer.

3.1.13. Cis makebae Souza-Gonçalves et Lopes-Andrade, 2017

Cis makebae Souza-Gonçalves & Lopes-Andrade 2017: 341 [11]. Type locality: Republic of South Africa: Limpopo Province, Sekororo Kloof.

Diagnosis: The species belongs to the neserorum group. It differs from other southern African species of the nesesorum group (except for C. afer, C. aster, C. bicaesariatus sp. n., C. caffer and C. masekelai) in males being devoid of concave impression in anterior pronotal portion. Cis makebae differs from C. afer, C. aster, C. bicaesariatus sp. n. and C. caffer in males bearing comparatively shorter anterocephalic and pronotal plates.

Host fungi:Trametes cingulata Berk. (Polyporaceae), two breeding records (Souza-Gonçalves & Lopes-Andrade 2017).

Distribution. Ethiopian. Known from northeastern KwaZulu-Natal and southeastern Limpopo (Republic of South Africa).

Comments: The species is known only from the type series [11]. This species was collected together with C. grobbelaarae sp. n., C. makrosoma sp. n., C. mandelai and the parasitoid A. micans [11].

3.1.14. Cis makrosoma Souza-Gonçalves et Lopes-Andrade, sp. n.

ZooBank: http://www.zoobank.org/NomenclaturalActs/BD37E9B0-CD68-4768-A7B9-5926792596CA

Figure 6(A–H); Figure 13.

Cis sp. G and Cis sp. O in Neser [9].

Type locality: “Nelspruit”, coordinates 25°29’ S 30°59’ E (Mbombela Municipality, Ehlanzeni District, Mpumalanga Province).

Etymology: The species name derives from the Greek adjective “makros”, which means “long”, and the Greek noun “soma”, which means “body”, both in the genitive singular. The name is a reference to the body shape of this species.

Diagnosis: The species belongs to the makrosoma group. It differs from all other southern African species (except for C. grobbelaarae sp. n.) by the pattern of the dual elytral punctation, consisting of megapunctures forming more or less regular longitudinal rows, in-between rows filled with micropunctures bearing short bristles. It differs from C. grobbelaarae sp. n. in males with anterocephalic edge truncate, a very elongated and flattened body, and moderately long and flattened prosternum.

Description, male holotype (Figure 6(A–D)): Adult fully pigmented and in good conditions, except for lacking the left antenna and four legs. Measurements in mm: TL 1.88, PL 0.64, PW 0.74, EL 1.24, EW 0.78, GD 0.52. Ratios: PL/PW 0.87, EL/EW 1.59, EL/PL 1.92, GD/EW 0.66, TL/EW 2.41. Body very elongate, flattened, dorsum and venter reddish dark brown; antennae, palpi and tarsi yellowish brown; dorsal vestiture single, consisting of short suberect bristles, easily discernible in low magnifications (<50×); ventral vestiture of decumbent setae, easily discernible in low magnifications (<50×). Head with anteriormost portion visible from above; dorsum with coarse and deep punctures, separated from each other by one puncture-width or less, with decumbent minute bristle (0.01–0.02 mm) arising from each puncture; interspaces, microreticulate; anterocephalic edge truncate. Antennae with 10 antennomeres, lengths as follows (in mm, right antennae measured): 0.05, 0.05, 0.04, 0.03, 0.02, 0.02, 0.02, 0.04, 0.04, 0.05 (FL 0.13 mm, CL 0.12 mm, CL/FL 0.95). Eyes coarsely facetted, with about 60 ommatidia; GW 0.15 mm. Gula 0.44× as wide as head. Pronotum (Figure 6(D)) with coarse, deep, single punctation, bearing an impunctate median line beginning around four puncture-widths of base until disc; punctures distributed irregularly, separated from each other by one to two puncture-widths; interspaces, microreticulate; vestiture single, consisting of suberect yellowish short bristles (~0.01 mm); anterior edge rounded; lateral edges crenulate, barely explanate and completely visible when seen from above; anterior corners barely angulate. Scutellar shield subtriangular, bearing few punctures and few bristles; BW 0.12 mm; SL 0.06 mm. Elytra with seriate, dual punctation; megapunctures coarse, deep, about 2× as large as micropunctures, separated from each other by one megapuncture-width or less, forming more or less longitudinal rows, in-between rows filled with micropunctures; interspaces, smooth and shiny; vestiture single, consisting of short suberect yellowish bristles (~0.01 mm) arising from micropunctures. Metathoracic wings developed, apparently functional. Hypomera with coarse, shallow punctation; each puncture bearing one fine decumbent seta; interspaces, microreticulate. Prosternum in front of coxae flattened, moderately long; interspaces, microreticulate. Prosternal process subparallel-sided, about 0.9× as long as prosternum at midline, apex rounded. Protibiae with maximum width about one-third of its length; apical edge devoid of spines; outer apical angle projected in acute tooth. Meso- and metatibiae without spines in apical edge. Metaventrite with coarse, deep punctures; interspaces, microreticulate; discrimen about one-third the length of metaventrite at midline. Abdominal ventrites with coarse, moderately deep punctures, separated from each other by one puncture-width or less and bearing one fine decumbent yellowish seta; interspaces, microreticulate; length of ventrites (in mm, from base to apex at the longitudinal midline) as follows: 0.33, 0.11, 0.10. 0.08, 0.11; first abdominal ventrite bearing a margined, large, circular, setose sex patch at center, with transverse diameter of 0.07 mm. Male terminalia in a paratype (Figure 6(F–H)) with sternite VIII (laid in bad position during dissection and not shown in figures) with posterior margin almost straight, bearing short setae at middle and long at rounded corners; anterior portion membranous. Tegmen (Figure 6(G)) 2.0× as long as wide, widest at apical third; sides expanding to apical third; one shallow emargination in each side at apex, forming acute angulation (Figure 6(G), small black arrows) and with rounded corners (Figure 6(G), big black arrows); anterior portion subtriangular. Basal piece (Figure 6(F)) semicircular, 1.6× as wide as long. Penis (Figure 6(H)) 0.6× as long as tegmen, 3.0× as long as wide; subparallel-sided; apex rounded and with one excavation in each side (Figure 6(H), red arrows); shortly rounded emargination at anterior portion.

Females (Figure 6(E)): Tenerals. Anterior edge of head truncate and anterior edge of pronotum rounded. Otherwise like males, but devoid of abdominal sex patch and protibial tooth.

Variation: Males, measurements in mm (n = 4, including the holotype): TL 1.89–1.92 (1.77 ± 0.21), PL 0.49–0.66 (0.59 ± 0.08), PW 0.57–0.74 (0.68 ± 0.08), EL 0.96–1.27 (1.17 ± 0.14), EW 0.68–0.79 (0.75 ± 0.05), GD 0.41–0.54 (0.50 ± 0.06). Ratios: PL/PW 0.86–0.89 (0.87 ± 0.01), EL/EW 1.42–1.66 (1.57 ± 0.10), EL/PL 1.92–2.08 (1.98 ± 0.07), GD/EW 0.61–0.73 (0.67 ± 0.05), TL/EW 2.15–2.46 (2.36 ± 0.15). Females, measurements in mm (n = 3): TL 1.55–1.73 (1.63 ± 0.09), PL 0.53–0.68 (0.59 ± 0.08), PW 0.60–0.65 (0.63 ± 0.03), EL 0.98–1.10 (1.04 ± 0.06), EW 0.65–0.68 (0.67 ± 0.01), GD 0.43–0.48 (0.45 ± 0.03). Ratios: PL/PW 0.81–1.08 (0.95 ± 0.14), EL/EW 1.50–1.63 (1.56 ± 0.07), EL/PL 1.56–2.10 (1.78 ± 0.20), GD/EW 0.65–0.70 (0.67 ± 0.03), TL/EW 2.38–2.56 (2.45 ± 0.09).

Type material: Holotype: ♂ (SANC) “SOUTH AFRICA: MPU, Nelspruit, 25°29’ S 30°59’ E, 14.viii.2009, D. van Heerden\Ex bracket fungus, #178, on Acacia sieberiana var. woodii\Ex bracket fungus Trametes sp., BF# 178\NATIONAL COLL. OF INSECTS Pretoria, S. Afr.\Cis makrosoma Souza-Gonçalves & Lopes-Andrade HOLOTYPUS [red paper]”. Paratypes: 3 ♂♂ and 3 ♀♀ as follows: 2 ♂♂ (1 CELC, dissected; 1 SANC) same data as the holotype; 1 ♂ (SANC) “SOUTH AFRICA: LIMP, Sekororo Kloof nr. Penge, 24°25’ S 30°27’ E, 27.vii.2008, S. Neser\Ex bracket fungus Trametes cingulata, BF# 155\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 3 ♀♀ (1 CELC; 2 SANC) “N.K. Wildtuin, G.A. Hepburn, 24.3.1960, Ac.X.950\NATIONAL COLL. OF INSECTS Pretoria, S. Afr.”. All paratypes additionally labeled “Cis makrosoma Souza-Gonçalves & Lopes-Andrade PARATYPUS [yellow paper]”.

Host fungi:Trametes sp., one record; Trametes cingulata Berk. (Polyporaceae), one record.

Distribution: Ethiopian. Known from eastern Mpumalanga and southeastern Limpopo (Republic of South Africa) (Figure 13).

Comments: This species was collected with C. makebae, C. mandelai, C. urbanae, the invasive species Cer. tabellifer and the parasitoid A. micans.

3.1.15. Cis mandelai Souza-Gonçalves et Lopes-Andrade, 2017

Cis mandelai Souza-Gonçalves & Lopes-Andrade 2017: 344 [11]. Type locality: Republic of South Africa: North Western Province, Marethwane.

Diagnosis: The species belongs to the neserorum group. It differs from other southern African species of the neserorum group (except for C. neserorum, C. stalsi, C. testaceus, and C. urbanae) in males bearing a concave impression in anterior pronotal portion. Cis mandelai differs from C. neserorum and C. stalsi by its prosternal process, which is conspicuously narrow near the base and gradually expanded to a rounded apex. It differs from C. urbanae in the less robust body and males with comparatively shorter pronotal plates; and from C. testaceus in bearing head covered by pronotum when seen from above and comparatively shorter pronotal plates with straight sides.

Host fungi:Ganoderma sp. (Ganodermataceae), one breeding record. Trametes sp. one breeding record; and Trametes cingulate Berk., one breeding record [11].

Distribution: Ethiopian. Known from eastern North West, western Gauteng, eastern Mpumalanga and northern Limpopo (Republic of South Africa).

Comments: The species is known only from the type series [11]. It was collected together with C. bicaesariatus sp. n., C. grobelaarae sp. n., C. makebae, C. paraliacus, C. parvisetosus sp. n., C. neserorum, C. westerncapensis sp. n., the morphospecies Cis sp. Y and Orthocis sp. A, the invasive species Cer. tabellifer and the parasitoid A. micans [11].

3.1.16. Cis masekelai Souza-Gonçalves et Lopes-Andrade, 2017

Cis masekelai Souza-Gonçalves & Lopes-Andrade 2017: 347 [11]. Type locality: Republic of South Africa: Mpumalanga Province, Die Hel Nature Reserve.

Diagnosis: The species belongs to the neserorum group. It differs from other southern African species of the neserorum group (except for C. afer, C. aster, C. bicaesariatus sp. n., C. caffer and C. makebae) in males being devoid of concave impression in anterior pronotal portion. Cis masekelai differs from C. afer, C. aster, C. bicaesariatus sp. n., C. caffer and C. makebae in males with anterior pronotal edge with a shallow emargination forming two very close short projections.

Host fungi:Trametes meyenii (Klotzsch) Lloyd, two record, one being a breeding record [11].

Distribution: Ethiopian. Known from eastern North West, northern Limpopo and eastern Mpumalanga (Republic of South Africa) [11].

Comments: The species is known only from the type series [10]. The records provided here (Figure 15) are corrections for that provided by Souza-Gonçalves and Lopes-Andrade [11]. In that paper, the authors cited one recorded as 25°45’ S 27°49’ E instead of 23°45’ S 27°49’ E (lapsus calami), here represented by the northernmost record (Figure 15). This species was collected together with C. neserorum and the invasive species Cer. tabellifer.

3.1.17. Cis mpumalangaensis Souza-Gonçalves et Lopes-Andrade, sp. n.

ZooBank: http://www.zoobank.org/NomenclaturalActs/80860930-35E0-49E8-8B4E-6167E984E971

Figure 7(A–I); Figure 13.

Cis sp. Q and Cis sp. N (in part) in Neser [9].

Type locality: “Mooihoek Farm” (near Wakkerstroom), coordinates 27°13’ S 30°32’ E (Pixley Ka Seme Local Municipality, Gert Sibande District, Mpumalanga Province).

Etymology: The species name is Latinized from “Mpumalanga” in the genitive singular. The name is a reference to the Mpumalanga Province, the province where the holotype and most part of the paratypes were collected.

Diagnosis: The species belongs to the pacificus group. It differs from C. foveocephalus sp. n. in males being devoid of a vertexal sex patch; from C. parvisetosus sp. n. in the comparatively thicker and denser vestiture, as well as longer and more acute anterocephalic plates; and from C. umlalaziensis sp. n. in the non-subseriate elytral vestiture.

Description, male holotype (Figure 7(A–D)): Adult fully pigmented and in good conditions, except for lacking one tarsus. Measurements in mm: TL 1.47, PL 0.49, PW 0.63, EL 0.98, EW 0.69, GD 0.56. Ratios: PL/PW 0.78, EL/EW 1.42, EL/PL 2.01, GD/EW 0.81, TL/EW 2.13. Body elongate, convex, dorsum and venter reddish dark brown; antennae, palpi and tarsi yellowish brown; dorsal vestiture single, consisting of short suberect bristles, easily discernible in high magnifications (>65×); ventral vestiture of decumbent setae easily discernible in high magnifications (>65×). Head with anteriormost portion visible from above; dorsum with coarse, deep punctures, separated from each other by less than one puncture-width, with decumbent minute bristles (0.01–0.02 mm) arising from each puncture; interspaces, microreticulate; anterocephalic edge produced and elevated forming two subtriangular plates. Antennae with 10 antennomeres, lengths as follows (in mm, left antennae measured): 0.05, 0.04, 0.04, 0.03, 0.02, 0.01, 0.01, 0.03, 0.04, 0.05 (FL 0.11 mm, CL 0.12 mm, CL/FL 1.14). Eyes coarsely facetted; with about 50 ommatidia; GW 0.13 mm. Gula 0.46× as wide as head. Pronotum (Figure 7(D)) with irregularly distributed, dual punctation, bearing an impunctate median line beginning about four puncture-widths of base until disc; megapunctures coarse, deep, about 2× as large as micropunctures, separated from each other by one megapuncture-width or less; interspaces, microreticulate; vestiture single, consisting of short suberect yellowish bristles (0.01–0.02 mm) arising from megapunctures; anterior edge rounded; lateral edges not crenulate, not explanate and not visible when seen from above; anterior corners rounded. Scutellar shield pentagonal, bearing few punctures and few bristles; BW 0.10 mm; SL 0.05 mm. Elytra with non-seriate, dual punctation; megapunctures coarse, deep, about 2× as large as micropunctures, separated from each other by two puncture-widths or less; interspaces a bit rugose; vestiture single, consisting of short suberect yellowish bristles (0.01–0.02 mm) arising from megapunctures. Metathoracic wings developed, apparently functional. Hypomera with coarse, shallow punctation; each puncture bearing one fine decumbent seta; interspaces, microreticulate. Prosternum in front of coxae biconcave and barely carinate; interspaces, microreticulate. Prosternal process subparallel-sided, about 0.8× as long as prosternum at midline, apex rounded. Protibiae with maximum width about one-third of its length; apical edge devoid of spines; outer apical angle projected in acute tooth. Meso- and metatibiae without spines in apical edge. Metaventrite with coarse, deep punctures; interspaces, microreticulate; discrimen about one-fourth the length of metaventrite at midline. Abdominal ventrites with coarse, moderately deep punctures, separated from each other by one puncture-width or less and bearing one fine decumbent yellowish seta; interspaces, microreticulate; length of ventrites (in mm, from base to apex at the longitudinal midline) as follows: 0.22, 0.08, 0.07, 0.07, 0.08; first abdominal ventrite bearing a margined, circular, setose sex patch at middle, with transverse diameter of 0.05 mm. Male terminalia in a paratype (Figure 7(F–I)) with sternite VIII (Figure 7(F)) with posterior margin almost straight, bearing short setae at middle and long setae at rounded corners; anterior portion membranous. Tegmen (Figure 7(H)) 1.9× as long as wide, widest at apex; sides expanding from basal third to apex; apex with deep V-shaped emargination forming slender lateral struts curved to middle (Figure 7(H), black arrows). Basal piece (Figure 7(G)) subtriangular, 1.4× as wide as long. Penis (Figure 7(I)) as long as tegmen, 4.6 as long as wide; subcylindrical, subparallel-sided and converging to triangular apex; anterior portion rounded.

Females (Figure 7(E)): Anterior edge of head truncate and anterior edge of pronotum rounded. Otherwise like males, but devoid of head plates, abdominal sex patch and protibial tooth.

Variation: Males, measurements in mm (n = 5, including the holotype): TL 1.20–1.53 (1.41 ± 0.13), PL 0.40–0.49 (0.47 ± 0.04), PW 0.50–0.63 (0.58 ± 0.05), EL 0.80–1.05 (0.95 ± 0.09), EW 0.60–0.73 (0.67 ± 0.05), GD 0.50–0.56 (0.53 ± 0.02). Ratios: PL/PW 0.78–0.83 (0.81 ± 0.02), EL/EW 1.33–1.46 (1.41 ± 0.05), EL/PL 2.00–2.10 (2.02 ± 0.04), GD/EW 0.76–0.83 (0.80 ± 0.03), TL/EW 2.00–2.19 (2.11 ± 0.07). Females, measurements in mm (n = 8): TL 1.13–1.45 (1.37 ± 0.12), PL 0.38–0.48 (0.43 ± 0.04), PW 0.45–0.63 (0.54 ± 0.06), EL 0.75–1.00 (0.93 ± 0.08), EW 0.55–0.70 (0.64 ± 0.05), GD 0.43–0.55 (0.51 ± 0.04). Ratios: PL/PW 0.75–0.90 (0.81 ± 0.05), EL/EW 1.36–1.56 (1.45 ± 0.06), EL/PL 2.00–2.33 (2.15 ± 0.13), GD/EW 0.68–0.84 (0.80 ± 0.05), TL/EW 2.05–2.28 (2.13 ± 0.08).

Type material: Holotype: ♂ (SANC) “SOUTH AFRICA: MPU, Mooihoek Farm, nr. Wakkerstroom, 27°13’ S 30°32’ E, 15.vii.2008, O & S Neser\Ex bracket fungus Coriolus versicolor, BF# 135\NATIONAL COLL. OF INSECTS Pretoria, South Africa\Cis mpumalangaensis Souza-Gonçalves & Lopes-Andrade HOLOTYPUS [red paper]”. Paratypes: 40 ♂♂ and 32 ♀♀ as follows: 13 ♂♂ (4 CELC, one dissected; 9 SANC) and 15 ♀♀ (5 CELC; 10 SANC) same data as the holotype; 11 ♂♂ (4 CELC, one dissected; 7 SANC) and 5 ♀♀ (3 CELC, one dissected; 2 SANC) “SOUTH AFRICA: MPU, Mooihoek Farm, nr. Wakkerstroom, 27°13’ S 30°32’ E, 15.vii.2008, O & S Neser\Ex bracket fungus Thelephora sp., BF# 125\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♂ and 2 ♀♀ (CELC) “SOUTH AFRICA: MPU, Mooihoek Farm, nr. Wakkerstroom, 27°13’ S 30°32’ E, 15.vii.2008, O & S Neser\Ex bracket fungus BF# 107\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 4 ♂♂ (2 CELC; 2 SANC) and 8 ♀♀ (3 CELC; 5 SANC) “SOUTH AFRICA: MPU, Mooihoek Farm, nr. Wakkerstroom, 27°13’ S 30°32’ E, 15.vii.2008, O & S Neser\Ex bracket fungus Coriolus versicolor, BF# 140\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 7 ♂♂ (3 CELC; 4 SANC) and 6 ♀♀ (2 CELC; 4 SANC) “SOUTH AFRICA: MPU, Mooihoek Farm, nr. Wakkerstroom, 27°13’ S 30°32’ E, 15.vii.2008, O & S Neser\Ex bracket fungus Stereum ostrea, BF# 138\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 2 ♂♂ (CELC) “SOUTH AFRICA: MPU, Mooihoek Farm, nr. Wakkerstroom, 27°13’ S 30°32’ E, 15.vii.2008, O & S Neser\Ex bracket fungus Stereum ostrea, BF# 90\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♂ (SANC) “SOUTH AFRICA: MPU, Mooihoek Farm, nr. Wakkerstroom, 27°13’ S 30°32’ E, 15.vii.2008, O & S Neser\Ex bracket unidentified fungus, BF# 168\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 2 ♀♀ (CELC) “SOUTH AFRICA: MPU, Mooihoek Farm, nr. Wakkerstroom, 27°13’ S 30°32’ E, 15.vii.2008, O & S Neser\Ex bracket fungus Trametes sp., BF# 113\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♂ (SANC) “SOUTH AFRICA: MPU, Mooihoek Farm, nr. Wakkerstroom, 27°13’ S 30°32’ E, 15.vii.2008, O & S Neser\Ex bracket fungus Ganoderma applanatum, BF# 139\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 2 ♀♀ (SANC) “SOUTH AFRICA: LIMP, Blouberg Mt. NW Polokwane, 23°04’ S 29°00’ E, 27.iv.2007, OC Neser\Adults ex bracket fungus on fallen trunk\Ex bracket fungus Trametes sp., BF# 34\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♀ (SANC) “SOUTH AFRICA: LIMP, Otter’ S Dan 16 Km from Hoedspruit, 24°24’ S 30°49’ E, 18.vii.2008, D. van Heerden\Ex bracket fungus Coriolus versicolor, BF# 136\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♀ (SANC) “SOUTH AFRICA: MPU, Alkmaar W Nelspruit, 25°27’ S 30°50’ E, 10.ii.2008\OC Neser\Ex bracket fungus Trametes versicolor, BF# 45\NATIONAL COLL. OF INSECTS Pretoria, South Africa”. All paratypes additionally labeled “Cis mpumalangaensis Souza-Gonçalves & Lopes-Andrade PARATYPUS [yellow paper]”.

Host fungi:Ganoderma applanatum (Pers.) Pat., one record; Stereum ostrea (Blume & T. Nees) Fr., two records, one being breeding record; Thelephora sp. (Thelephoraceae), one breeding record; Trametes sp., two records (obs.: we are not sure whether these correspond to a single fungus species or several unidentified Trametes); Trametes versicolor (L.) Lloyd, four records, two being breeding records.

Distribution: Ethiopian. Known from eastern and southeastern Mpumalanga, northern and southeastern Limpopo (Republic of South Africa) (Figure 13).

Comments: This species was collected with C. bicaesariatus sp. n., C. foveocephalus sp. n., C. neseroum, C. mooihoekite, C. tessariplacus sp. n., Xylographus madagascariensis Mellié, 1849, Scydmaeninae sp., the invasive species Cer. tabellifer, and the parasitoids A. gracilis Neser, 2012, A. micans and A. silvani Neser, 2012.

3.1.18. Cis mooihoekite Souza-Gonçalves et Lopes-Andrade, 2018

Cis mooihoekite Souza-Gonçalves & Lopes-Andrade 2018: 27 [12]. Type locality: Republic of South Africa: Mpumalanga Province, Mooihoek Farm.

Diagnosis: The species belongs to the bilamellatus group. It differs from C. pickeri in the TL less than 1.30 mm; pronotum devoid of a median impunctate line; anterocephalic edge in male with acute corners and pronotal plate angularly emarginate forming two small and triangular corners with acute apex; and male abdominal sex patch about one-quarter the length of the first ventrite at midline.

Host fungi:Thelephora sp., one record; and Trametes versicolor (L.) Lloyd, two records [12].

Distribution: Ethiopian. Known from northern and southern Mpumalanga (Republic of South Africa).

Comments: The species is known only from the type series [12]. This species was collected together with C. grobbelaarae sp. n., X. madagascariensis, the invasive species Cer. tabellifer and the parasitoids A. micans and A. silvani.

3.1.19. Cis muriceus Mellié, 1849

Cis muriceus Mellié 1849: 348 [22]. Type locality: Republic of South Africa: Western Cape Province, Cape of Good Hope.

Host fungi. Unknown.

Distribution: Ethiopian. Known from Cape of Good Hope (Western Cape Province, Republic of South Africa).

Comments: The species is known only from the type series [22]. There is no further record in the literature, as far as we have traced. We cannot place it in any group at this moment.

3.1.20. Cis neserorum Souza-Gonçalves et Lopes-Andrade, 2017

Cis neserorum Souza-Gonçalves & Lopes-Andrade 2017: 349 [11]. Type locality: Republic of South Africa: Limpopo Province, Wesfalia Estate.

Diagnosis: The species belongs to the neserorum group. It differs from other southern African species of the neserorum group (except for C. mandelai, C. stalsi, C. testaceus, and C. urbanae) in males bearing a concave impression in anterior pronotal portion. Cis neserorum differs from C. mandelai, C. stalsi and C. urbanae in males with first abdominal ventrite devoid of a sex patch. It differs from C. testaceus in possessing comparatively shorter and closer pronotal plates.

Additional material: 3 ♂♂ and 3 ♀♀ as follows: 1 ♂ (ANIC) “REP. SOUTH AFRICA: Natal, 75 Km WSW Estcourt Cathedral Peaks For. Sta., 7-31.XII.74, S. & J. Peck\Ber 28, 29.XII.79, Podocarp forest, rotted wood w/ fungi, 1500m”; 2 ♂♂ and 2 ♀♀ (ANIC) “NATAL, A. JANSE\J.F. Lawrence Lot.2008\ex Polyporus sanguineus\ex U.S.D.A. Herbaria”; 1 ♀ (CUIC) “Port St. Johns, U. S. AFR., Mar.6-8.1949, J.C. Bradley\CU [salmon paper]”. All additionally labeled “Cis neserorum Souza-Gonçalves & Lopes-Andrade, 2017, I. Souza-Gonçalves det.”

Host fungi:Ganoderma applanatum (Pers.) Pat., one record; Lenzites elegans (Spreng.) Pat. (Polyporaceae), six breeding records; Pycnoporus sanguineus (L.) Murrill (Polyporaceae), ten records, eight being breeding records; Stereum ostrea (Blume & T. Nees) Fr. (Steraceae), one breeding record; Trametes sp., six records, three being breeding records (obs.: we are not sure whether these correspond to a single fungus species or several unidentified Trametes); Trametes hirsuta (Wulfen) Lloyd, four records, three being breeding records; Trametes meyenii (Klotzsch) Lloyd, two breeding records; Trametes polyzona (Pers.) Justo, four records, three of which are breeding records; Trametes versicolor (L.) Lloyd, seven records, six being breeding records; Thelephora sp., one breeding record [11].

Distribution: Ethiopian. Known from many localities in Republic of South Africa [11]. The species is the largest distributed among Cis species in southern and South Africa [11]. The additional material was collected in western and southern KwaZulu-Natal (Republic of South Africa) (Figure 15).

Comments: This species was collected together with C. chinensis, C. grobbelaarae sp. n., C. fuscipes, C. mandelai, C. masekelai, C. mooihoekite, C. mpumalangaensis sp. n., C. parvisetosus sp. n., C. pickeri, C. stalsi, C. westerncapensis sp. n., the morphospecies Cis sp. S, Cis sp. Y, Orthocis sp. A, X. madagascariensis, the invasive species Cer. tabellifer and the parasitoids A. micans, A. gracilis and A. silvani.

3.1.21. Cis paraliacus Souza-Gonçalves et Lopes-Andrade, 2018

Cis paraliacus Souza-Gonçalves & Lopes-Andrade 2018: 510 [13]. Type locality: Republic of South Africa: KwaZulu-Natal Province, Umlalazi Nature Reserve.

Diagnosis: The species belongs to the multidentatus group. It differs from C. chinensis in bearing elytral punctation dual and lateral protonal edges not visible from above.

Host fungi:Trametes hirsuta (Wulfen) Lloyd, one breeding record [13].

Distribution: Ethiopian. Known from eastern and southeastern KwaZulu-Natal (Republic of South Africa).

Comments: The species is known only from the type series [13]. This species was collected together with C. mandelai, C. umlalaziensis sp. n., the invasive species Cer. tabellifer, the tenebrionid Pentaphyllus fronticornis Gebien, 1910, and the parasitoid A. micans [13].

3.1.22. Cis parvisetosus Souza-Gonçalves et Lopes-Andrade, sp. n.

ZooBank: http://www.zoobank.org/NomenclaturalActs/33ED4C68-7474-4316-BB3C-6734A1067C85

Figure 8(A–I); Figure 9(A–J) Figure 14.

Cis sp. M and Cis sp. N in Neser [9].

Type locality: “Monk’ S Cowl Foothills” (Champagne Castle), coordinates 29°03’ S 29°23’ E (Drakensberg range, KwaZulu-Natal Province).

Etymology: The species name derives from the Latin adjectives “parvus”, which means “small”, and “setosum”, which means “setose”, both in the genitive singular. The name is a reference to the short vestiture of this species.

Diagnosis: The species belongs to the pacificus group. It differs from C. foveocephalus sp. n. in males being devoid of a vertexal sex patch; from C. mpumalangaensis sp. n. in the comparatively thinner and sparser vestiture, and comparatively shorter and less acute anterocephalic plates; and from C. umlalaziensis sp. n. in the non-subseriate and comparatively thinner elytral vestiture.

Description, male holotype (Figure 8(A–D)): Adult fully pigmented and in good conditions, except for lacking two tarsi and both antennae. Measurements in mm: TL 1.34, PL 0.46, PW 0.58, EL 0.88, EW 0.65, GD 0.51. Ratios: PL/PW 0.80, EL/EW 1.35, EL/PL 1.89, GD/EW 0.79, TL/EW 2.07. Body elongate, convex, dorsum and venter reddish dark brown; palpi and tarsi yellowish brown; dorsal vestiture single, consisting of minute suberect setae, easily discernible in high magnifications (>80×); ventral vestiture of decumbent setae easily discernible in high magnifications (>80×). Head with anteriormost portion visible from above; dorsum with coarse and deep punctures, separated from each other by one puncture-width or less, with suberect minute seta (0.01–0.02 mm) arising from each puncture; interspaces, microreticulate; anterocephalic produced and slightly elevated forming two subtriangular. Antennae with 10 antennomeres, lengths as follows (in mm, left antennae measured in a paratype): 0.06, 0.05, 0.03, 0.04, 0.03, 0.02, 0.01, 0.03, 0.03, 0.06 (FL 0.13 mm, CL 0.12 mm, CL/FL 0.95). Eyes coarsely facetted, with about 60 ommatidia; GW 0.13 mm. Gula 0.55× as wide as head. Pronotum (Figure 8(D)) with irregularly distributed, dual punctation, bearing an impunctate median line beginning five puncture-widths of base until disc; megapunctures coarse, deep, about 2× as large as micropunctures, separated from each other by one megapuncture-width or less; interspaces, microreticulate; vestiture single, consisting of minute suberect pale yellowish setae (0.01–0.02 mm) arising from megapunctures; anterior edge rounded; lateral edges not crenulate, not explanate and not visible when seen from above; anterior corners rounded. Scutellar shield triangular, bearing few punctures and apparently glabrous; BW 0.09 mm; SL 0.08 mm. Elytra with non-seriate, dual punctation; megapunctures coarse, deep, about 2× as large as micropunctures, separated from each other by two megapuncture-widths or less; interspaces a bit rugose; vestiture single, consisting of minute suberect pale yellowish setae (0.01–0.02 mm) arising from megapunctures. Metathoracic wings developed, apparently functional. Hypomera with coarse, deep punctation; each puncture bearing a fine decumbent seta; interspaces, microreticulate. Prosternum in front of coxae biconcave and barely carinate; interspaces, microreticulate. Prosternal process subparallel-sided, about 0.9× as long as prosternum at midline, apex rounded. Protibiae with maximum width about one-fourth of its length; apical edge devoid of spines; outer apical angle rounded. Meso- and metatibiae without spines in apical edge. Metaventrite with coarse, deep punctures; interspaces, microreticulate; discrimen about one-fifth the length of metaventrite at midline. Abdominal ventrites with coarse, moderately deep punctures, separated from each other by one puncture-width or less and bearing one fine decumbent pale yellowish seta; interspaces, microreticulate; length of ventrites (in mm, from base to apex at the longitudinal midline) as follows: 0.23, 0.08, 0.08, 0.08, 0.09; first abdominal ventrite bearing margined, circular, setose sex patch at middle, with transverse diameter of 0.05 mm. Male terminalia in a paratype (Figure 8(F–I)) with sternite VIII (Figure 8(F)) with posterior margin almost straight, bearing short setae at middle and long setae at rounded corners. Tegmen (Figure 8(H)) 1.4× as long as wide, widest at apex; sides expanding from basal third to apex; apex with shallow V-shaped emargination; corners rounded (Figure 8(H), black arrows); anterior portion subtriangular. Basal piece (Figure 8(G)) subhexagonal, 2.8× as long as wide. Penis (Figure 8(I)) 1.1× as long as tegmen, 4.0× as long as wide; subcylindrical, subparallel-sided and converging to subtriangular apex; anterior portion rounded.

Females (Figure 8(E)): Anterior edge of head truncate and anterior edge of pronotum rounded. Otherwise like males, but devoid of abdominal sex patch.

Variation: Males, measurements in mm (n = 9, including the holotype): TL 0.91–1.50 (1.26 ± 0.17), PL 0.31–0.50 (0.42 ± 0.06), PW 0.38–0.60 (0.52 ± 0.07), EL 0.60–1.00 (0.84 ± 0.12), EW 0.43–0.70 (0.59 ± 0.08), GD 0.33–0.53 (0.46 ± 0.06). Ratios: PL/PW 0.79–0.90 (0.82 ± 0.03), EL/EW 1.35–1.56 (1.43 ± 0.06), EL/PL 1.84–2.06 (1.98 ± 0.07), GD/EW 0.65–0.90 (0.79 ± 0.07), TL/EW 2.07–2.33 (2.15 ± 0.08). In some populations, the cephalic plates are smaller than than those of males from the type locality (Figure 9(F)) or absent (Figure 9(A)). Some differences are also noted in aedeagus from different populations, principally in size of tegmen and shape of basal piece (Figure 9(B–E,G–J)). Females, measurements in mm (n = 16): TL 0.85–1.53 (1.23 ± 0.17), PL 0.30–0.48 (0.41 ± 0.05), PW 0.30–0.48 (0.41 ± 0.05), EL 0.35–0.58 (0.49 ± 0.06), EW 0.55–1.05 (0.82 ± 0.12), GD 0.40–0.70 (0.57 ± 0.08). Ratios: PL/PW 0.75–0.94 (0.84 ± 0.05), EL/EW 1.31–1.52 (1.43 ± 0.06), EL/PL 1.83–2.27 (2.02 ± 0.11), GD/EW 0.65–0.86 (0.79 ± 0.05), TL/EW 1.96–2.26 (2.15 ± 0.08).

Type material: Holotype: ♂ (SANC) “SOUTH AFRICA: KZN, Monk’ S Cowl Foothills Drakensberg, 29°03’ S 29°23’ E, 24.iii.2008, S. & O.C. Neser\Ex bracket fungus on fallen log, BF# 64\Ex bracket fungus Hymenochaete ochromarginata, BF# 64\NATIONAL COLL. OF INSECTS Pretoria, South Africa\Cis parvisetosus Souza-Gonçalves & Lopes-Andrade HOLOTYPUS [red paper]”. Paratypes: 8 ♂♂ and 20 ♀♀ as follows: 1 ♂ (CELC, dissected) and 2 ♀♀ (1 CELC; 1 SANC) same data as the holotype; 1 ♂ and 2 ♀♀ (SANC) “SOUTH AFRICA: WCAPE, Garden of Eden Indig. Forest, nr. Knysna, 34°02’ S 23°12’ E, 1.iii.1991, AJ Hendricks\Emerged from log of Nuxia floribunda BUDDLEJACEAE, UA677B\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♂ and 1 ♀ (CELC) “SOUTH AFRICA: WCAPE, Groenkop Indigenous For., near George, 33°56’ S 22°31’ E, 22.iii.1991, AJ Hendricks\Emerged from dead branches of living tree of Olinia ventosa OLINIACEAE, UA703\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♀ (CELC) “SOUTH AFRICA: WCAPE, Groenkop Indigenous For., near George, 33°56’ S 22°31’ E, 25.iii.1991, AJ Hendricks\Emerged from dead branches of living tree Rhus chiruadensis ANACARDIACEAE, UA717\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♂ (SANC) “SOUTH AFRICA: WCAPE, Heatherlands, George, 33°57’ S 22°27’ E, 11.iv.1990, AJ Urban\Emerged from Dais cotinifolia THYMELAEACEAE, with fruiting body of fungus ?Lenzites, UA601\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♂ (CELC, dissected) and 4 ♀♀ (2 CELC; 2 SANC) “SOUTH AFRICA: WCAPE, Prince Alfred’ S Pass N of Knysna, 33°58’ S 23°09’ E, 5.xi.2009, S & OC Neser\Ex bracket fungus #215 on fallen tree trunk\Ex bracket fungus Fomitopsis lilacino-gilva, BF# 215\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♂ and 1 ♀ (CELC) “SOUTH AFRICA WCAPE, Prince Alfred’ S Pass N of Knysna, 33°58’ S 23°09’ E, 5.xi.2009, S & OC Neser\Ex bracket fungus Coriolus versicolor, BF# 123\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 3 ♀♀ (SANC) “SOUTH AFRICA WCAPE, Montagu Pass N of George, 33°54’ S 22°24’ E, 4.xi.2009, S & OC Neser\Ex bracket fungus, #208 on fallen tree trunk\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 2 ♂♂ (1 CELC, dissected; 1 SANC) and 3 ♀♀ (1 CELC; 2 SANC) “SOUTH AFRICA: KZN, Mpisini Nature Res., 30°12’ S 30°48’ E, 9.vii.2008, S. & O.C. Neser\Ex bracket fungus Funalia sp., BF# 110\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♀ (SANC) “SOUTH AFRICA: WCAPE, Montagu Pass N of George, 33°54’ S 22°24’ E, 4.xi.2009, S. & O.C. Neser\Ex bracket fungus, #201 on fallen tree trunk\Ex bracket fungus Ganoderma applanatum, BF# 201\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 2 ♀♀ (CELC) “SOUTH AFRICA: WCAPE, Montagu Pass N of George, 33°54’ S 22°24’ E, 4.xi.2009, S. & O.C. Neser\Ex bracket fungus, #210 on fallen tree trunk\Ex bracket fungus Coriolus hirsutus, BF# 210\NATIONAL COLL. OF INSECTS Pretoria, South Africa”. All paratypes additionally labeled “Cis parvisetosus Souza-Gonçalves & Lopes-Andrade PARATYPUS [yellow paper]”.

Host fungi:Funalia sp. (Polyporaceae), one record; Ganoderma applanatum (Pers.) Pat., one record; Hymenochaete ochromarginata P.H.B. Talbot (Hymenochaetaceae), one record; Trametes hirsuta (Wulfen) Lloyd, one record; Trametes versicolor (L.) Lloyd, one record; Rhodofomitopsis lilacinogilva (Berk.) B.K. Cui, M.L. Han & Y.C. Dai, one record.

Distribution: Ethiopian. Known from southern Western Cape, southeastern and western KwaZulu-Natal (Republic of South Africa) (Figure 14).

Comments: This may be a cryptic species and further studies are necessary to elucidate this case. It was collected with C. fuscipes, C. neserorum, C. pickeri, C. regius, C. westerncapensis sp. n., the morphospecies Cis sp. S, Orthocis sp. A, Ptinidae sp. 3, X. madagascariensis, the invasive species Cer. tabellifer, and the parasitoids A. gracilis and A. naiadis.

3.1.23. Cis pickeri Lopes-Andrade, Matushkina, Buder et Klass, 2009

Cis pickeri Lopes-Andrade et al. 2009:57 [1]. Type locality: Republic of South Africa: Western Cape Province, Ceres; Souza-Gonçalves & Lopes-Andrade 2018: 31 [12] {new distributional records from Republic of South Africa}.

Diagnosis: The species belongs to the bilamellatus group. It differs from C. mooihoekite in the TL being more than 1.30 mm; pronotum with a median impunctate line; anterocephalic edge in male with rounded corners and pronotal plate slightly emarginate forming two small projections with subrounded apex; and male abdominal sex patch about one-third the length of the first ventrite at midline.

Host fungi:Laetiporus sp. (Fomitopsidaceae), one record; Russula capensis A. Person (Russulaceae), one record; and Trametes versicolor (L.) Lloyd, one breeding record [12].

Distribution: Ethiopian. Known from southern Western Cape (Republic of South Africa).

Comments: The species was collected together with C. bicaesariatus sp. n., C. grobbelaarae sp. n., C. chinensis, C. parvisetosus sp. n., C. neserorum, the morphospecies Cis sp. S, Cis sp. Y and the invasive species Cer. tabellifer [12].

3.1.24. Cis regius Orsetti et Lopes-Andrade, 2016

Cis regius Orsetti & Lopes-Andrade 2016: 146 [10]. Type locality: Republic of South Africa: Western Cape Province, Prince Alfred’ S Pass.

Diagnosis: The species belongs to the regius group. It differs from all southern African Cis species by the peculiar occipital tubercle close to vertex in males; pronotum with dual punctation, lateral to anterior edges broadly rounded and bearing a row of scattered setae; and elytra with single punctation and vestiture of seriate setae.

Additional material: 5 ♂♂ and 4 ♀♀ as follows: 2 ♂♂ (1 CELC, dissected; 1 SANC) and 1 ♀ (SANC) “SOUTH AFRICA: WCAPE, Storms River Mouth, Tsitsikamma National Park, 34°01’ S 23°54’ E, 29.v.2003, S. Neser\Ex bracket fungus [E]\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♀ (SANC) “SOUTH AFRICA WCAPE, Diepwalle State Foret, 33°58’ S 23°09’ E, 1.iii.1990, AJ Urban & AJ Hendricks\Collected from seedlings in shaded nursery of Psoralea pinnata FABACEAE, UA590:A\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♂ and 1 ♀ (ANIC) “REP. S. AFRICA: Cape Prov., Knysna, Diepwalle, 450 m, 17 Dec. 1981, 81–170, forest fungi & litter, S. & J. Peck, coll.”; 2 ♂♂ and 1 ♀ (ANIC) “REP. S. AFRICA: Cape Prov., Gouna, 81–186, 23 Dec. 1981, berlese forest log litter, S. & J. Peck, coll.”. All additionally labeled “Cis regius Orsetti & Lopes-Andrade, 2016; I. Souza-Gonçalves det.”.

Host fungi:Ganoderma applanatum (Pers.) Pat., one breeding record [10].

Distribution: Ethiopian. Known from southern Western Cape (Republic of South Africa). The additional material was collected in four areas near the type locality (Figure 15).

Comments: This species was collected together with C. parvisetosus sp. n. and the invasive species Cer. tabellifer.

3.1.25. Cis stalsi Souza-Gonçalves et Lopes-Andrade, 2017

Cis stalsi Souza-Gonçalves & Lopes-Andrade 2017: 352 [11]. Type locality: Republic of South Africa: KwaZulu-Natal Province, Hlabisi.

Diagnosis: The species belongs to the neserorum group. It differs from other southern African species of the neserorum group (except for C. mandelai, C. neserorum, C. testaceus, and C. urbanae) in males bearing a concave impression in anterior pronotal portion. Cis stalsi differs from C. neserorum in males with first abdominal ventrite bearing a sex patch. It differs from C. mandelai and C. urbanae in possessing subparallel-sided prosternal process with rounded apex; and from C. testaceus in the comparatively shorter pronotal plates and very close anterocephalic plates.

Additional material: 3 ♂♂ and 1 ♀ as follows: 1 ♂♂ and 1 ♀ (ANIC) “REP. SOUTH AFRICA: Natal, 75 Km WSW Estcourt Cathedral Peaks For. Sta., 7-31.XII.79, S. & J. Peck\Ber 22, 26.XII.79, Eucalyptus logs, frass & fungi & decaying bark, 1400m”; 2 ♂♂ (ANIC) “REP. SOUTH AFRICA: Natal, 75 km WSW Estcourt Cathedral Peaks For. Sta., 7-31.XII.79, S. & J. Peck\Ber 23, 26.XII.79, Eucalyptus logs, decayed litter & soil, 1400 m”. All additionally labeled “Cis stalsi Souza-Gonçalves & Lopes-Andrade, 2017; I. Souza-Gonçalves det.”

Host fungi:Podoscypha parvula (Lloyd) D. A. Reid (Meruliaceae), one breeding record [11].

Distribution: Ethiopian. Known from eastern North West and eastern KwaZulu-Natal (Republic of South Africa). The additional material was collected in western KwaZulu-Natal (Republic of South Africa) (Figure 15).

Comments: This species was collected together with C. neserorum and the invasive species Cer. tabellifer [11].

3.1.26. Cis tessariplacus Souza-Gonçalves et Lopes-Andrade, sp. n.

ZooBank: http://www.zoobank.org/NomenclaturalActs/C9186CD7-27CB-4551-BF4B-8E256E3F810C

Figure 10(A–I); Figure 14.

Cis sp. L in Neser [9].

Type locality: “Mooihoek Farm” (near Wakkerstroom), coordinates 27°13’ S 30°32’ E (Pixley Ka Seme Local Municipality, Gert Sibande District, Mpumalanga Province).

Etymology: The species name derives from the Greek noun “tessares”, which means “four”, and “plakos”, which means “plates”, both in the genitive singular. The name is a reference to the number of plates present on the anterocephalic edge of this species.

Diagnosis: The species resembles members of the Cis multidentatus group in the anterocephalic edge produced and elevate forming four teeth in males and anterior pronotal edge in male projected into two triangular plates; but the prosternum is biconcave and the outer apical angle of protibia is projected in an acute tooth only in males.

Description, male holotype (Figure 10(A–D)): Adult apparently not fully pigmented but in good conditions, except for lacking one leg, four tarsi, the left antenna, and for being covered by dust or fungus. Measurements in mm: TL 1.22, PL 0.50, PW 0.75, EL 1.22, EW 0.77, GD 0.59. Ratios: PL/PW 0.67, EL/EW 1.58, EL/PL 2.44, GD/EW 0.77, TL/EW 2.23. Body elongate, convex, dorsum reddish brown (except for dust-covered areas, but visible elsewhere in paratypes); venter dark reddish brown; antenna yellowish brown with club dark brown, palpi and tarsi yellowish brown; dorsal vestiture dual, consisting of short suberect bristles and minute decumbent setae, easily discernible in high magnifications (>65×); ventral vestiture of decumbent setae easily discernible in high magnifications (>65×). Head with anteriormost portion visible from above; dorsum with coarse and deep punctures, separated from each other by less than one puncture-width, with one minute decumbent bristle (0.01–0.02 mm) arising from each puncture; interspaces, microreticulate; anterocephalic edge produced and elevated forming four acute and triangular teeth. Antennae with 10 antennomeres, lengths as follows (in mm, right antennae measured): 0.05, 0.05, 0.05, 0.03, 0.02, 0.02, 0.01, 0.04, 0.04, 0.06 (FL 0.13 mm, CL 0.14 mm, CL/FL 1.07). Eyes coarsely facetted, with about 60 ommatidia; GW 0.14 mm. Gula 0.56× as wide as head. Pronotum (Figure 10(D)) with coarse, deep, single punctation, devoid of an impunctate median line; punctures distributed irregularly, separated from each other by less one puncture-width; interspaces, microreticulate; vestiture single, consisting of short suberect bristles (0.03–0.04 mm); anterior edge produced and elevated forming two triangular plates with acute apex; lateral edges barely crenulate, not explanate and not visible when seen from above; anterior corners rounded. Scutellar shield pentagonal, bearing few punctures and few bristles; BW 0.13 mm; SL 0.07 mm. Elytra with non-seriate, single punctation; punctures coarse, deep, distributed irregularly, separated from each other by two puncture-widths or less; interspaces, somewhat a bit rugose; vestiture dual, consisting of short suberect bristles (~0.04 mm) and minute decumbent setae (~0.02 mm), both yellowish and arising from punctures. Metathoracic wings developed, apparently functional. Hypomera with coarse, shallow punctation; each puncture bearing one fine decumbent seta; interspaces, microreticulate. Prosternum in front of coxae biconcave; interspaces, microreticulate. Prosternal process subparallel-sided, as long as prosternum at midline, apex rounded. Protibiae with maximum width about one-third of its length; apical edge devoid of spines; outer apical angle projected in acute tooth. Meso- and metatibiae without spines in apical edge. Metaventrite with coarse, deep punctures; interspaces, microreticulate; discrimen about two-fifths the length of metaventrite at midline. Abdominal ventrites with coarse, moderately deep punctures, separated from each other by one puncture-width or less and bearing one fine decumbent yellowish seta; interspaces, microreticulate; length of ventrites (in mm, from base to apex at the longitudinal midline) as follows: 0.24, 0.10, 0.08, 0.07, 0.09; first abdominal ventrite bearing a margined, small, circular, setose sex patch at middle, with transverse diameter of 0.03 mm. Male terminalia in a paratype (Figure 10(F–I)) with sternite VIII (Figure 10(F)) with posterior margin deeply emarginate, bearing short setae at middle and long setae at acute corners; anterior margin membranous. Tegmen (Figure 10(H)) 2.7× as long as wide, widest at middle; small emargination at apex, globular tubercles at apical portion (Figure 10(H), black arrows). Basal piece (Figure 10(G)) subtrapezoidal, 1.4× as wide as long. Penis (Figure 10(I)) as long as tegmen, 4.2× as long as wide; sides expanding until apical third and then converging to apex; apical portion membranous and bearing acute sclerotization at middle (Figure 10(I), big red arrow); truncate sclerotization at anterior portion (Figure 10(I), small red arrows).

Females (Figure 10(E)): Anterior edge of head truncate and anterior edge of pronotum rounded. Otherwise like males, but devoid of abdominal sex patch and protibial tooth.

Variation: Males, measurements in mm (n = 2, including the holotype): TL 1.55–1.72 (1.62 ± 0.14), PL 0.50–0.53 (0.51 ± 0.02), PW 0.60–0.75 (0.67 ± 0.10), EL 1.00–1.22 (1.11 ± 0.16), EW 0.65–0.77 (0.71 ± 0.09), GD 0.48–0.59 (0.53 ± 0.08). Ratios: PL/PW 0.67–0.88 (0.77 ± 0.14), EL/EW 1.54–1.58 (1.56 ± 0.03), EL/PL 1.90–2.44 (2.17 ± 0.38), GD/EW 0.73–0.77 (0.75 ± 0.03), TL/EW 2.23–2.39 (2.29 ± 0.08). Female, measurements in mm (n = 1): TL 1.73, PL 0.58, PW 0.68, EL 1.15, EW 0.83, GD 0.60. Ratios: PL/PW 0.85, EL/EW 1.39, EL/PL 2.00, GD/EW 0.73, TL/EW 2.09.

Type material: Holotype: ♂ (SANC, dissected) “SOUTH AFRICA: MPU, Mooihoek Farm, nr. Wakkerstroom, 27°13’ S 30°32’ E, 15.vii.2008, O & S Neser\Ex bracket fungus Stereum ostrea, BF# 138\ NATIONAL COLL. OF INSCTS Pretoria, South Africa\Cis tessariplacus Souza-Gonçalves & Lopes-Andrade HOLOTYPUS [red paper]”. Paratypes: 1♂ and 1 ♀ as follows: 1♀ (SANC) same data as the holotype; 1♂ (CELC, dissected) “SOUTH AFRICA: MPU, Alkmaar W Nelspruit, 25°27’ S 30°50’ E, 10.ii.2008\OC Neser\Ex bracket fungus Trametes sp., BF# 47\NATIONAL COLL. OF INSECTS Pretoria, South Africa”. All paratypes additionally labeled “Cis tessariplacus Souza-Gonçalves & Lopes-Andrade PARATYPUS [yellow paper]”.

Host fungi:Stereum ostrea (Blume & T. Nees) Fr., one record; Trametes sp., one record.

Distribution: Ethiopian. Known from eastern and southeastern Mpumalanga (Republic of South Africa) (Figure 14).

Comments: This species was collected together with C. foveocephalus sp. n., C. mpumalangaensis sp. n., the invasive species Cer. tabellifer, and the parasitoids A. gracilis and A. naiadis. We prefer not to place it in any group at this moment.

3.1.27. Cis testaceus Fåhraeus, 1871

Cis testaceus Fåhraeus, 1871: 671 [21]. Type locality: Caffraria (=Kaffraria), currently Republic of South Africa: Eastern Cape Province (no specific locality); Ferrer 1997: 408 [24] (lectotype designation).

Host fungi: Unknown.

Distribution: Ethiopian. Known from Eastern Cape Province (Republic of South Africa).

Comments: The species is known only from the type series [21]. There is no further record in the literature, as far as we have traced. It is a member of the neserorum group.

3.1.28. Cis urbanae Souza-Gonçalves et Lopes-Andrade, 2017

Cis urbanae Souza-Gonçalves & Lopes-Andrade 2017: 354 [11]. Type locality: Republic of South Africa: Mpumalanga Province, Nelspruit.

Diagnosis: The species belongs to the neserorum group. It differs from other southern African species of the neserorum group (except for C. mandelai, C. neserorum, C. stalsi and C. testaceus) in males bearing a concave impression in anterior pronotal portion. Cis urbanae differs from C. neserorum in males with first abdominal ventrite bearing a sex patch, and from C. neserorum and C. stalsi in the prosternal process conspicuously narrow near base and gradually expanding to a rounded apex. Cis mandelai bears a similar prosternal process, but differs from in the comparatively more robust body and males with comparatively longer pronotal plates. It differs from C. testaceus in the more robust body.

Host fungi:Trametes sp., one breeding record [11].

Distribution: Ethiopian. Known from eastern Mpumalanga (Republic of South Africa).

Comments: The species is known only from the type series [11]. It was collected together with C. makrosoma sp. n., the invasive species Cer. tabellifer and the parasitoids A. micans.

3.1.29. Cis umlalaziensis Souza-Gonçalves et Lopes-Andrade, sp. n.

ZooBank: http://www.zoobank.org/NomenclaturalActs/2D0198D9-6915-441E-81A5-FA15FFBCE815

Figure 11(A–G); Figure 13.

Cis sp. K in Neser [9].

Type locality: “Umlalazi Nature Reserve” (near Mtunzini), coordinates 28°57’ S 31°46’ E (uMlalazi Local Municipality, Uthungulu District, KwaZulu-Natal Province).

Etymology: The species name is Latinized from “Umlalazi” in the genitive singular. The name is a reference to the Umlalazi Nature Reserve, the type locality of this species.

Diagnosis: The species belongs to the pacificus group. It differs from C. foveocephalus sp. n. in males being devoid of vertexal sex patch; from C. mpumalangaensis sp. n. in the subseriate elytral vestiture; and from C. parvisetosus sp. n. in the comparatively thicker and subseriate vestiture, and comparatively longer and more acute anterocephalic plates.

Description, male holotype (Figure 11(A–D)): Adult fully pigmented in good conditions, except for lacking both antennae. Measurements in mm: TL 1.11, PL 0.39, PW 0.49, EL 0.72, EW 0.50, GD 0.41. Ratios: PL/PW 0.79, EL/EW 1.45, EL/PL 1.87, GD/EW 0.83, TL/EW 2.22. Body elongate, convex, dorsum and venter reddish dark brown; palpi and tarsi yellowish brown; dorsal vestiture single, consisting of short suberect bristles, easily discernible in high magnifications (>95×); ventral vestiture of decumbent setae easily discernible in high magnifications (>95×). Head with anteriormost portion visible from above; dorsum with coarse and deep punctures, separated from each other by one puncture-width or less, with minute decumbent seta (~0.01 mm) arising from each puncture; interspaces, microreticulate; anterocephalic edge produced and elevated forming two triangular plates. Antennae with 10 antennomeres, lengths as follows (in mm, right antennae measured in a paratype): 0.05, 0.04, 0.03, 0.02, 0.01, 0.01, 0.01, 0.02, 0.03, 0.05 (FL 0.08 mm, CL 0.10 mm, CL/FL 1.25). Eyes coarsely facetted, with about 50 ommatidia; GW 0.11 mm. Gula 0.48× as wide as head. Pronotum (Figure 11(D)) with irregularly distributed, dual punctation, bearing an impunctate median line beginning around four puncture-widths of base until disc; megapunctures coarse, deep, about 2× as large as micropunctures, separated from each other by one puncture-width or less; interspaces, microreticulate; vestiture single, consisting of short suberect yellowish bristles (0.01–0.02 mm) arising from megapunctures; anterior edge rounded; lateral edges not crenulate, not explanate and not visible when seen from above; anterior corners rounded. Scutellar shield pentagonal, bearing few punctures and few bristles; BW 0.05 mm; SL 0.04 mm. Elytra with non-seriate, dual punctation; megapunctures coarse, deep, about 2× as large as micropunctures, separated from each other by two puncture-widths or less; interspaces a bit rugose; vestiture subseriate, single, consisting of short suberect yellowish bristles (0.01–0.02 mm) arising from megapunctures. Metathoracic wings developed, apparently functional. Hypomera with coarse, shallow punctation; each puncture bearing one fine decumbent seta; interspaces, microreticulate. Prosternum in front of coxae biconcave and barely carinate; interspaces, microreticulate. Prosternal process subparallel-sided, about as long as prosternum at midline, apex rounded. Protibiae with maximum width about one-third of its length; apical edge devoid of spines; outer apical angle projected in acute tooth. Meso- and metatibiae without spines in apical edge. Metaventrite with coarse, deep punctures; interspaces, microreticulate; discrimen about one-third the length of metaventrite at midline. Abdominal ventrites with coarse, moderately deep punctures, separated from each other by one puncture-width or less and bearing one fine decumbent yellowish seta; interspaces, microreticulate; length of ventrites (in mm, from base to apex at the longitudinal midline) as follows: 0.15, 0.06, 0.05, 0.05, 0.07; first abdominal ventrite bearing a unmargined, small, oval, setose sex patch at middle, with transverse diameter of 0.02 mm. Male terminalia in a paratype (Figure 11(E–G)) with sternite VIII (Figure 11(E)) with posterior margin almost straight, bearing short setae at middle and long setae at subrounded corners; anterior margin membranous. Tegmen (Figure 11(F)) 2.8× as long as wide, widest at apical third; sides expanding to apex; apex with one emargination in each side forming three rounded lobes (Figure 11(F), black arrows), the lateral ones slightly shorter than the mid one; anterior portion rounded. Penis (Figure 11(G)) 1.2× as long as tegmen, 5.8× as long as wide; subcylindrical, subparallel-sided and converging to subtriangular apex; anterior portion rounded.

Female: Unknown.

Variation: Males, measurements in mm (n = 3, including the holotype): TL 0.98–1.11 (1.05 ± 0.07), PL 0.38–0.39 (0.38 ± 0.01), PW 0.45–0.49 (0.48 ± 0.02), EL 0.60–0.72 (0.68 ± 0.06), EW 0.48–0.50 (0.50 ± 0.01), GD 0.40–0.43 (0.40 ± 0.01). Ratios: PL/PW 0.79–0.83 (0.79 ± 0.03), EL/EW 1.26–1.45 (1.35 ± 0.09), EL/PL 1.60–1.87 (1.80 ± 0.14), GD/EW 0.80–0.89 (0.80 ± 0.05), TL/EW 2.05–2.22 (2.10 ± 0.09).

Type material: Holotype: ♂ (SANC) “SOUTH AFRICA: KZN, Umlalazi Nature Res., Mtunzini, 28°57’ S 31°46’ E, 13.vii.2008, R.P. Urban\Ex bracket fungus Coriolus hirsutus, BF# 87\NATIONAL COLL. OF INSECTS Pretoria, South Africa\Cis umlalaziensis Souza-Gonçalves & Lopes-Andrade HOLOTYPUS [red paper]”. Paratypes: 2 ♂♂ (1 CELC, dissected; 1 SANC) same data as the holotype. All paratypes additionally labeled “Cis umlalaziensis Souza-Gonçalves & Lopes-Andrade PARATYPUS [yellow paper]”.

Host fungi:Trametes hirsuta (Wulfen) Lloyd, one record.

Distribution: Ethiopian. Known from eastern KwaZulu-Natal (Figure 13).

Comments: This species was collected together with C. paraliacus, the invasive species Cer. tabellifer, the tenebrionid P. fronticornis and the parasitoid A. micans.

3.1.30. Cis westerncapensis Souza-Gonçalves et Lopes-Andrade, sp. n.

ZooBank:http://www.zoobank.org/NomenclaturalActs/0DC523DE-0682-4E23-A051-28AC566ADE91

Figure 12(A–I).

Cis sp. J in Neser [9].

Type locality: “Montagu Pass” coordinates 33°54’ S 22°24’ E (north of George, Western Cape Province).

Etymology: The species name is Latinized from “Western Cape” in the genitive singular. The name is a reference to the Western Cape Province, the unique known province where the species occurs.

Diagnosis: The species belongs to the westerncapensis group. It differs from C. lacinipennis sp. n. in bearing slightly longer (0.03–0.04 mm) and subseriate vestiture, lighter dorsal coloration and a comparatively wider body.

Description, male holotype (Figure 12(A–D)): Fully pigmented and in good conditions, except for lacking the right antenna and one tarsus. Measurements in mm: TL 1.47, PL 0.49, PW 0.58, EL 0.98, EW 0.69, GD 0.55. Ratios: PL/PW 0.84, EL/EW 1.42, EL/PL 2.01, GD/EW 0.80, TL/EW 2.13. Body elongate, convex, dorsum and venter reddish brown; antenna yellowish brown with club dark brown, palpi and tarsi yellowish brown; dorsal vestiture single, consisting of short suberect bristles, easily discernible in high magnifications (>75×); ventral vestiture of decumbent setae easily discernible in high magnifications (>75×). Head with anteriormost portion visible from above; dorsum with coarse and deep punctures, separated from each other by one punctures-width or less, with minute decumbent bristle (0.01–0.02 mm) arising from each puncture; interspaces, microreticulate; anterocephalic edge truncate. Antennae with 10 antennomeres, lengths as follows (in mm, left antenna measured): 0.06, 0.04, 0.03, 0.02, 0.02, 0.02, 0.02, 0.04, 0.04, 0.06 (FL 0.10 mm, CL 0.14 mm, CL/FL 1.42). Eyes coarsely facetted, with about 50 ommatidia; GW 0.12 mm. Gula 0.46× as wide as head. Pronotum (Figure 12(D)) with irregularly distributed, dual punctation, devoid of impunctate median line; megapunctures coarse, deep, about 2× as large as micropunctures, separated from each other by one megapuncture-width or less; interspaces microreticulate; vestiture single, consisting of moderately short suberect pale yellowish bristles (0.02–0.03 mm) arising from micropunctures; anterior edge rounded; lateral edges not crenulate, not explanate and not visible when seen from above; anterior corners rounded. Scutellar shield pentagonal, bearing few punctures and apparently glabrous; BW 0.08 mm; SL 0.05 mm. Elytra with subseriate, dual punctation; megapunctures coarse, deep, about 2× as large as micropunctures, separated from each other by two megapuncture-widths or less; interspaces a bit rugose; vestiture subseriate, single, consisting of moderately short suberect pale yellowish bristles (0.03–0.04 mm) arising from megapunctures. Metathoracic wings developed, apparently functional. Hypomera with coarse, shallow punctation; each puncture bearing one fine decumbent seta; interspaces, microreticulate. Prosternum in front of coxae biconcave and barely carinate; interspaces, microreticulate. Prosternal process subparallel-sided, about 0.9× as long as prosternum at midline, apex rounded. Protibiae with maximum width about one-fourth of its length; apical edge devoid of spines; outer apical angle projected in acute tooth. Meso- and metatibiae without spines in apical edge. Metaventrite with coarse, deep punctures; interspaces, microreticulate; discrimen about one-third the length of metaventrite. Abdominal ventrites with coarse, moderately deep punctures, separated from each other by one puncture-width or less and bearing one fine decumbent yellowish seta; interspaces, microreticulate; length of ventrites (in mm, from base to apex at the longitudinal midline) as follows: 0.22, 0.08, 0.08, 0.08, 0.10; first abdominal ventrite bearing one margined, large, circular, setose sex patch at middle, with transverse diameter of 0.07 mm. Male terminalia in a paratype (Figure 12(F–I)) with sternite VIII (Figure 12(F)) with posterior margin almost straight, bearing short setae at middle and long setae at rounded corners; anterior portion membranous. Tegmen (Figure 12(H)) 1.8× as long as wide, widest at apex; subparallel-sided; two acute angulations in each side at apex (Figure 12(H), black arrows); anterior portion subrounded. Basal piece (Figure 12(G)) subtriangular, 1.4× as wide as long. Penis (Figure 12(I)) 4.3× as long as tegmen, 10.0× as long as wide; subcylindrical, subparallel-sided, somewhat stick-shaped; anterior and posterior portions rounded.

Females (Figure 12(E)): Anterior edge of head truncate and anterior edge of pronotum rounded. Otherwise like males, but devoid of abdominal sex patch and protibial tooth.

Variation: Males, measurements in mm (n = 21, including the holotype): TL 1.23–1.65 (1.35 ± 0.10), PL 0.43–0.58 (0.46 ± 0.04), PW 0.48–0.65 (0.33 ± 0.05), EL 0.80–1.08 (0.90 ± 0.07), EW 0.55–0.80 (0.63 ± 0.06), GD 0.45–0.63 (0.51 ± 0.04). Ratios: PL/PW 0.77–0.95 (0.86 ± 0.05), EL/EW 1.33–1.58 (1.43 ± 0.70), EL/PL 1.75–2.24 (1.97 ± 0.13), GD/EW 0.78–0.88 (0.81 ± 0.02), TL/EW 2.04–2.33 (2.16 ± 0.09). Females, measurements in mm (n = 28): TL 1.23–1.58 (1.36 ± 0.09), PL 0.40–0.53 (0.45 ± 0.04), PW 0.45–0.58 (0.51 ± 0.03), EL 0.83–1.05 (0.91 ± 0.06), EW 0.53–0.70 (0.62 ± 0.04), GD 0.45–0.58 (0.51 ± 0.04). Ratios: PL/PW 0.81–1.00 (0.89 ± 0.05), EL/EW 1.36–1.58 (1.46 ± 0.06), EL/PL 1.81–2.24 (2.00 ± 0.12), GD/EW 0.76–0.90 (0.83 ± 0.04), TL/EW 2.07–2.38 (2.20 ± 0.08).

Type material: Holotype: ♂ (SANC) “SOUTH AFRICA WCAPE, Montagu Pass N of George, 33°54’ S 22°24’ E, 4.xi.2009, S & OC Neser\Ex bracket fungus, #210 on fallen tree trunk\Ex bracket fungus Coriolus hirsutus, BF# 210\NATIONAL COLL. OF INSECTS Pretoria, South Africa\ Cis westerncapensis Souza-Gonçalves & Lopes-Andrade HOLOTYPUS [red paper]”. Paratypes: 39 ♂♂ and 56 ♀♀ as follows: 1 ♀ (SANC) same data as the holotype; 2 ♀♀ (SANC) “SOUTH AFRICA WCAPE, Montagu Pass N of George, 33°54’ S 22°24’ E, 4.xi.2009, S & OC Neser\Ex bracket fungus, #211 on Brachylaena neriifolia\Ex bracket fungus Coriolus hirsutus, BF# 211”; 2 ♀♀ (CELC) “SOUTH AFRICA WCAPE, Montagu Pass N of George, 33°54’ S 22°24’ E, 4.xi.2009, S & OC Neser\Ex bracket fungus, #201 on fallen tree trunk\Ex bracket fungus Ganoderma applanatum, BF# 201\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♂ (CELC, dissected) and 1 ♀ (CELC) “SOUTH AFRICA WCAPE, Montagu Pass N of George, 33°54’ S 22°24’ E, 4.xi.2009, S & OC Neser\Ex bracket fungus, #199 on Rapanea sp.\Ex bracket fungus Ganoderma applanatum, BF# 199\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♂ and 1 ♀ (SANC) “SOUTH AFRICA, CP, Grootvadersbosch, nr. Heidelberg, 33.50S 20.53E, v.1985, A.J. Prins \ NATIONAL COLL. OF INSECTS Pretoria, S. Afr.”; 1 ♂ and 3 ♀♀ (CELC) “SOUTH AFRICA WCAPE, Prince Alfred’ S N of Knysna, 33°58’ S 23°09’ E, 5.xi.2009, S & OC Neser\Ex bracket fungus, #215 on fallen tree trunk\Ex bracket fungus Fomitopsis lilacino-gilva, BF# 215\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♀ (SANC) “SOUTH AFRICA WCAPE, Montagu Pass N of George, 33°54’ S 22°24’ E, 4.xi.2009, S & OC Neser\Ex bracket fungus Coriolus hirsutus, BF# 186\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♀ (SANC) “SOUTH AFRICA WCAPE, Montagu Pass N of George, 33°54’ S 22°24’ E, 4.xi.2009, S & OC Neser\Ex bracket fungus, #188 on fallen tree trunk\Ex bracket fungus Coriolus sp., BF# 188\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 3 ♂♂ (1 CELC, dissected; 2 SANC) and 3 ♀♀ (1 CELC, dissected; 2 SANC) “SOUTH AFRICA: WCape, Saasveld Foretry. Coll., 33°57’ S 22°32’ E, 14.viii.1990, AAJ Hendricks\Collected from rotten log of Olea capensis OLEACEAE, UA627\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 1 ♂ and 1 ♀ (SANC) “SOUTH AFRICA: WCape, Groeneweide Forest Walk, Saasveld, 33°58’ S 22°32’ E, 20.iii.1991, AJ Hendricks\Emerged from dead branches of living tree of Rapanea melanophloeos MYRSINIACEAE, UA716\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 2 ♂♂ (1 CELC; 1 SANC) “SOUTH AFRICA WCAPE, Prince Alfred’ S Pass N of Knysna, 35°58’ S 23°09’ E, 5.xi.2009, S & OC Neser\Ex bracket fungus, #219 on fallen tree trunk\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 24 ♂♂ (8 CELC; 16 SANC, 13 in dried capsule) and 31 ♀♀ (10 CELC; 21 SANC, 13 in dried capsule) “SOUTH AFRICA WCAPE, Montagu Pass N of George, 33°54’ S 22°24’ E, 4.xi.2009, S & OC Neser\Ex bracket fungus, #208 on fallen tree trunk\NATIONAL COLL. OF INSECTS Pretoria, South Africa”; 2 ♀♀ (SANC) “SOUTH AFRICA: WCAPE, Heatherlands, George, 33°57’ S 22°27’ E, 11.iv.1990, AJ Urban\Emerged from Dais cotinifolia THYMELAEACEAE, with fruiting body of fungus ?Lenzites, UA601”; 3 ♂♂ (1 CELC; 2 ANIC) and 5 ♀♀ (1 CELC; 4 ANIC) “REP. S. AFRICA: Cape Prov., Knysna, Diepwalle 450m, 17 Dec. 1981, 81-170, forest fungi & litter, S. & J. Peck, coll.”; 2 ♂♂ and 1 ♀ (ANIC) “REP. S. AFRICA: Cape Prov., Knysna, Gouna, 81-186, 23 Dec. 1981, berlese forest log litter, S. & J. Peck, coll.”; 1 ♂ and 1 ♀ (CELC) “REP. S. AFRICA: Cape Prov., Tsitsikama For. N.P., 20 Dec. 1981, 81-176, Berlese forest litter, S. & J. Peck, coll.”. All paratypes additionally labeled “Cis westerncapensis Souza-Gonçalves & Lopes-Andrade PARATYPUS [yellow paper]”.

Host fungi:Ganoderma applanatum (Pers.) Pat., two records; Rhodofomitopsis lilacinogilva (Berk.) (Berk.) B.K. Cui, M.L. Han & Y.C. Dai (Fomitopsidaceae), one record; Trametes sp., one record; Trametes hirsuta (Wulfen) Lloyd, three records.

Distribution: Ethiopian. Known from southern Western Cape (Figure 13).

Comments: This species was collected together with C. neserorum, C. parvisetosus sp. n., C. regius, the morphospecies Orthocis sp. A, X. madagascariensis, the invasive species Cer. tabellifer, and the parasitoids A. gracilis and A. naiadis.

3.2. Key to southern African species of Cis Latreille

Note: This identification key applies only to species described [1,9,10,11,12] or examined by us. The following southern Africa Cis species were not included: C. afer, C. bimucronatus, C. caffer, C. capensis, C. delagoensis, C. muriceus and C. testaceus. The key works for both males and females, except for C. fuscipes, of which only females are known to occur in southern Africa, and C. aster and C. umlalaziensis sp. n., of which only males are known.

1 Anterocephalic edge rounded, truncate, or projected in a single elevated plate or paired plates, horns, teeth or small angulations AND first abdominal ventrite with sex patch; IF sex patch absent, THEN both anterocephalic edge and anterior pronotal edge with short projections or subtriangular plates. Protibia with outer apical angle ALWAYS projected in an acute tooth. Males...........................................………………………………………………………………………………2

1’ Anterocephalic edge truncate or barely emarginate AND first abdominal ventrite without sex patch. Protibia with outer apical angle variable, simple, projected in an acute tooth or bearing sockted spines. Females……………………….. ……………………………………………………………………………….23

2 (1) Anterocephalic edge ALWAYS elevated and produced forming four teeth AND anterior pronotal edge projected in two small plates; IF pronotum with anterior edge simple, THEN head with occipital tubercle close to vertex….………………………………………….………………..………3

2’ Anterocephalic edge NEVER elevated and produced forming four teeth (can be elevated and produced forming two teeth, but NEVER four teeth). Pronotum with anterior edge variable……………………….. ……………………………………………………………………………….6

3 (2) Head with occipital tubercle close to vertex. Pronotum with anterior edge rounded; punctation dual. Elytral vestiture single, consisting of seriate setae. Tegmen with apex subtriangular delimited by internal excavations; penis with apex slightly enlarged and membranous. Southern WC……………………………………………….…….…….…….Cis regius Orsetti et Lopes-Andrade

3’ Head without occipital tubercle. Pronotum with anterior edge projected in two small plates; punctation single or inconspicuously dual. Elytral vestiture single, consisting of non-seriate bristles; IF vestiture dual, THEN consisting of mixed bristles and setae…………………………………………4

4 (3’) Prosternum biconcave. Elytral vestiture dual, consisting of mixed short bristles and minute setae. Tegmen with apex bearing small emargination and globular tubercles in apical portion (Figure 10(H); black arrows); penis with apical portion membranous and bearing acute sclerotization at middle (Figure 10(I); big red arrow), and base with truncate sclerotization (Figure 10(I); small red arrows). Eastern and southeastern MP…………………Cis tessariplacus Souza-Gonçalves et Lopes-Andrade sp. n.

4’ Prosternum almost flat to tumid at midline. Elytral vestiture single, consisting of non-suseriate bristles……...…………...…..…..…..…..…..…..…..…..…..…..…..…..…..…..…..…..…..…..…..…..…..…..5

5 (4’) Pronotum with punctures separated by less than one puncture-width; lateral margins barely to completely visible from above. Elytra with single punctation. Tegmen deeply and broadly emarginate at apex forming three lobes, the lateral ones barely acute and the mid one somewhat arrow-shaped; penis with basal third bearing lateral struts, which join to form membranous apex. Cosmopolitan; introduced at southern WC..………………………………..…..Cis chinensis Lawrence

5’ Pronotum with punctures separated by one to two puncture-widths; lateral margins not visible from above. Elytra with inconspicuously dual punctation. Tegmen deeply and broadly emarginate at apex forming three lobes, the lateral ones barely acute and the mid one somewhat arrow-shaped; penis with base bearing two short sclerotizations. Eastern and southeastern KZN….…………………………..…..…..…..……...Cis paraliacus Souza-Gonçalves et Lopes-Andrade

6 (2’) Elytral vestiture indistinctly to distinctly dual, IF indistinctly dual, THEN anterior pronotal edge barely to strongly emarginate forming short prominences or subtriangular plates.......................7

6’ Elytral vestiture single and anterior pronotal edge ALWAYS rounded….…….…..…..…….……...16

7 (6) Anterocephalic edge and anterior pronotal edge with one plate........…………..…..…..….....…….8

7’ Anterocephalic edge and anterior pronotal edge barely to strongly emarginate, forming two short prominences or subtriangular plates……………………………………..…..…..…..…..…..…..…..….…..9

8 (7) TL more than 1.30 mm. Anterocephalic edge with rounded corners. Pronotum with plate slightly emarginate forming two short projections with subrounded apex. Male with abdominal sex patch about one-third the length of first ventrite at midline. Tegmen bearing deep emargination at apex; penis subcylindrical and with apex bearing shallow emargination. Southern WC…………….................................................Cis pickeri Lopes-Andrade, Matushkina, Buder et Klass

8’ TL less than 1.30 mm. Anterocephalic edge with acute corners. Pronotum with plate angularly emarginate forming two small and triangular horns with acute apex. Male with abdominal sex patch about one-fourth the length of first ventrite at midline. Tegmen with apex bearing a small emargination on both sides; penis subcylindrical and with apex bearing short emargination. Northern and southern MP..…………..………...Cis mooihoekite Souza-Gonçalves et Lopes-Andrade

9 (7’) First male abdominal ventrite devoid of sex patch……………………………………..……...…...10

9’ First male abdominal ventrite bearing sex patch ………………...……………………..…..…..……...12

10 (9) Prosternal process relatively narrow, at least 4× as long as wide, no more than 0.15 as wide as gula. Tegmen elongate, with bilobed apex and membranous apical portion (Figure 8; black arrows); penis with three acute angulations at apex (Figure 9; red arrows). Northern MP, southeastern LP and northeastern NW…………………..…….Cis bicaesariatus Souza-Gonçalves et Lopes-Andrade sp. n.

10’ Prosternal process relatively wide, no more than 3× as long as wide, more than 0.20 as wide as gula……………………………………………........................…..…..…..…....…..…..…..…..…..…..….......11

11 (10’) Pronotum with anterior portion bearing concave impression; IF concave impression is inconspicuous, THEN head visible from above. Prosternal process with rounded apex. Tegmen elongate, devoid of lateral excavations or angulations, and apical portion with narrow emargination at middle; penis subcylindrical. EC, GP, KZN, LP, MP, NW and WC………………..…..Cis neserorum Souza-Gonçalves et Lopes-Andrade

11’ Pronotum with anterior portion devoid of concave impression AND head not visible from above. Prosternal process with rounded apex. Tegmen elongate, somewhat mace-shaped, with each lateral edge hardened near middle and bearing small concave emargination with acute angulation at base; penis very elongate………………………..……………...Cis aster Souza-Gonçalves et Lopes-Andrade

12 (9’) Prosternal process conspicuously narrow near base and gradually expanding to rounded apex………………………………………….………………………………..…..…..…..…..…..…..…..…...13

12’ Prosternal process subparallel-sided with rounded apex....………………..…..…..…..……………..14

13 (12) Body very robust. Lateral pronotal margins smooth. Male with first abdominal ventrite bearing glabrous sex patch. Tegmen elongate, devoid of lateral excavations or angulations, apical portion with membranous areas and rounded sclerotization; penis subcylindrical. Eastern MP………………..…..…..…..…..…..…....…..…….....Cis urbanae Souza-Gonçalves et Lopes-Andrade

13’ Body not conspicuously robust. Lateral pronotal margins weakly crenulate. Male with first abdominal ventrite bearing setose sex patch. Tegmen with three deep emarginations at apex; penis with apical portion membranous and with V-shaped sclerotization. Eastern NW, western GP, eastern MP and northern LP…………………….…Cis mandelai Souza-Gonçalves et Lopes-Andrade

14 (12’) Anterocephalic edge with two triangular plates separated from each other by less than half width of scutellar shield and with rounded apices. Pronotum with anterior portion bearing concave impression. Male with first abdominal ventrite with minute sex patch, about one-tenth the length of first ventrite at midline. Tegmen with apical portion emarginated forming two lateral lobes with rounded apices, inner edges close to each other; penis with struts not linked at base. Eastern NW, eastern and western KZN………………..…...…………Cis stalsi Souza-Gonçalves et Lopes-Andrade

14’ Anterocephalic edge with two triangular plates separated from each other by one width of scutellar shield or a bit less, and with subacute to acute apices. Pronotum with anterior portion devoid of concave impression. Male with first abdominal ventrite with sex patch at middle, more than one-fifth the length of first ventrite at midline……...…..…..…..…..…..…..…..…..…..…..…..…...15

15 (14’) Pronotum with anterior pronotal edge projected into two short triangular plates; apex, subacute; interspaces, barely microreticulate; anterior corners, rounded. Tegmen with apical portion bearing sclerotization at middle and laterals, which give the appearance of a trident; penis with two struts curved inwardly. Northeastern KZN and southeastern LP………………………….Cis makebae Souza-Gonçalves et Lopes-Andrade

15’ Pronotum with anterior edge emarginated anteriorly, forming two short projections; interspaces, coarsely microreticulate; anterior corners, angulate. Tegmen with membranous apical portion and bearing emargination at apex; penis with struts not linked at base and forming conspicuous sclerotizations, apex subtriangular and membranous. Eastern NW, northern LP and eastern MP………..………………………..……...…..…..….Cis masekelai Souza-Gonçalves et Lopes-Andrade

16 (6’) Elytra with punctation more or less to distinctly seriate………………………..…..…..………..17

16’ Elytra with punctation non-seriate…………………………………..……...………..…..…..………...20

17 (16) Lateral pronotal margins completely visible from above. Elytra with seriate and dual punctation; megapunctures forming more or less longitudinal rows, in-between rows filled with micropunctures; vestiture consisting of moderately short bristles arising from micropuntures.....…18

17’ Lateral pronotal margins not visible from above. Elytra with subseriate and dual punctation; megapunctures not forming longitudinal rows; vestiture consisting of moderately short bristles arising from megapunctures....……………………………………...…..…..…..…..…..…..…..…..…….....19

18 (17) Body very elongate and distinctly flattened. Anterocephalic edge truncate. Prosternum moderately long and flattened. Elytral vestiture consisting of short bristles (~0.01 mm). Tegmen with apex bearing one shallow emargination in each side and with acute angulations (Figure 6(G); big black arrows) and rounded corners (Figure 6(G); small black arrows); penis with rounded apex and with one excavation in each side (Figure 6(H); red arrows). Eastern MP and southeastern LP……Cis makrosoma Souza-Gonçalves et Lopes-Andrade sp. n.

18’ Body elongate and convex. Anterocephalic edge with small angulations. Prosternum concave. Elytral vestiture consisting of moderately short bristles (0.03–0.04 mm). Tegmen with apex bearing one deep emargination in each side and with acute angulations at corners (Figure 4(H); black arrows); penis with apical portion membranous and with a sclerotization at middle, apex truncate (Figure 4(I); red arrows). Northern MP and northwestern LP….…………….Cis grobbelaarae Souza-Gonçalves et Lopes-Andrade sp. n.

19 (17’) Color dark reddish brown. Elytral vestiture distinctly seriate, dense and short (0.02–0.03 mm). Male with first abdominal ventrite bearing unmargined sex patch. Tegmen with apex bearing rounded emargination and two small tubercles at middle (Figure 5(G); black arrows); penis with apex rounded and base with one flap in each side (Figure 5(H); red arrows). Western KZN and northern FS………………………..….….….….…….Cis lacinipennis Souza-Gonçalves et Lopes-Andrade sp. n.

19’ Color reddish brown. Elytral vestiture subseriate, sparse and long (0.03–0.04 mm). Male with first abdominal ventrite bearing margined sex patch. Tegmen with apex bearing two acute angulations in each side (Figure 12(H); black arrows); penis about 4× as long as tegmen and somewhat stick-shaped. Southern WC………………………....Cis westerncapensis Souza-Gonçalves et Lopes-Andrade sp. n.

20 (16’) Head with vertexal sex patch; anterocephalic edge with two small triangular tubercles. Tegmen with one rounded emargination in each side forming three lobes, the lateral ones short and acute (Figure 2(H); big black arrows), the mid one long and somewhat arrow-shaped (Figure 2(H); small black arrow); penis elongate and with membranous apex. Southeastern MP, northeastern NW and western KZN……………………………Cis foveocephalus Souza-Gonçalves et Lopes-Andrade sp. n.

20’ Head ALWAYS without vertexal sex patch; anterocephalic edge with two subtriangular or triangular plates……………………………………………...………….….….….….….….….….….….….21

21 (20’) Anterocephalic edges with plates longer than scutellar shield at midline. Elytral vestiture subseriate. Tegmen with apex bearing one emargination in each side, forming three rounded lobes (Figure 11(F); black arrows), the lateral ones slightly shorter than the mid one; penis subcylindrical and with subtriangular apex. Eastern KZN…………………………Cis umlalaziensis Souza-Gonçalves et Lopes-Andrade sp. n.

21’ Anterocephalic edges with plates shorter than scutellar shield at midline. Elytral vestiture non-subseriate..……………………………………………….….……..….….….….….….….….…...….…22

22 (21’) Dorsal vestiture consisting of thin and sparse setae. Tegmen apex bearing shallow somewhat V-shaped emargination and with rounded corners (Figure 9(B); black arrows); penis subcylindrical and with subtriangular apex. Southern WC, southeastern and western KZN…….....................................……….....Cis parvisetosus Souza-Gonçalves et Lopes-Andrade sp. n.

22’ Dorsal vestiture consisting of thick and dense bristles. Tegmen with apex bearing deep V-shaped emargination, forming lateral struts curved to middle (Figure 7(H); black arrows); penis subcylindrical and with triangular apex. Eastern and southeastern MP, northern and southeastern LP…………………….….….…….…Cis mpumalangaensis Souza-Gonçalves et Lopes-Andrade sp. n.

23 (1’) Elytra with punctation more or less to distinctly seriate…….............…….….….……………….24

23’ Elytra with punctation non-seriate………………………………….…………..….….…..…………...28

24 (23) Lateral pronotal margins completely visible from above. Elytral vestiture consisting of short or moderately long bristles arising from micropunctures….……….……...….….….….….….….…..…25

24’ Lateral pronotal margins not visible from above. Elytral vestiture consisting of short bristles arising from megapunctures……………………………….….….….….….…...….….….….….…………27

25 (24) Protibia with outer apical angle dentate. Cosmopolitan; introduced at southeastern MP and southern WC………………………………....….….….….….….….….….….……..…...Cis fuscipes Mellié

25’ Protibia with outer apical angle simple..……………………………….….…....….….….…………...26

26 (25’) Body very elongate and distinctly flattened. Prosternum moderately long and flattened. Elytral vestiture consisting of short bristles (~0.01 mm). Eastern MP and southeastern LP…...………………………………….…...Cis makrosoma Souza-Gonçalves et Lopes-Andrade sp. n.

26’ Body elongate and convex. Prosternum concave. Elytral vestiture consisting of moderately short bristles (0.03–0.04 mm). Northern MP and northwestern LP…………………………...Cis grobbelaarae Souza-Gonçalves et Lopes-Andrade sp. n.

27 (24’) Color dark reddish brown. Elytral vestiture dense, short (0.02–0.03 mm), distinctly seriate. Western KZN and northern FS….…….....Cis lacinipennis Souza-Gonçalves et Lopes-Andrade sp. n.

27’ Color reddish brown. Elytral vestiture subseriate, sparse and long (0.03–0.04 mm). Southern WC……………..…………..………….Cis westerncapensis Souza-Gonçalves et Lopes-Andrade sp. n.

28 (23’) Elytral vestiture distinctly dual; IF indistinctly dual, THEN punctation dual..………………29

28’ Elytral vestiture single……………………………………………………………...................................39

29 (28) Protibia with outer apical angle expanded and bearing socketed spines....................................30

29’ Protibia with outer apical angle not expanded and devoid of spines……………………...………..31

30 (29) TL more than 1.30 mm. Southern WC….……………..…………….................................Cis pickeri Lopes-Andrade Matushkina, Buder et Klass

30’ TL less than 1.30 mm. Northern and southern MP..………..…………………………Cis mooihoekite Souza-Gonçalves et Lopes-Andrade

31 (29’) Elytra with sigle punctation. Eastern and southeastern MP………….……...Cis tessariplacus Souza-Gonçalves et Lopes-Andrade sp. n.

31’ Elytra with dual punctation …………………………………………………………………………….32

32 (31’) Prosternal process conspicuously narrow near base and gradually expanding to rounded apex…..…………………………………………………………………………………………….………….33

32’ Prosternal process subparallel- or parallel-sided, with rounded or truncate apex….….………….35

33 (32); Body very robust. Lateral pronotal margins smooth Eastern MP…………………..Cis urbanae Souza-Gonçalves et Lopes-Andrade

33’ Body not conspicuosly robust. Lateral pronotal margins weakly crenulate. Eastern NW, western GP, eastern MP and northern LP…………………..Cis mandelai Souza-Gonçalves et Lopes-Andrade

35 (32’) Prosternal process relatively narrow, at least 4× as long as wide, no more than 0.15 as wide as gula. Northern MP, southeastern LP and northeastern NW…………..……………..Cis bicaesariatus Souza-Gonçalves & Lopes-Andrade sp. n.

35’ Prosternal process relatively wide, no more than 3× as long as wide, more than 0.20 as wide as gula………………………………………………………………………………………….............................36

36 (35’) Pronotal punctures usually separated by more than one puncture diameter and interspaces barely microreticulate. Northeastern KZN and southeastern LP.......……..………………..Cis makebae Souza-Gonçalves et Lopes-Andrade

36’ Pronotal punctures usually separated by less than one puncture-width and interspaces coarsely microreticulate………………………………………………………………………………………..………37

37 (36’) Pronotum with anterior corners angulate. Eastern NW, northern LP and eastern MP……………………..………………………………………Cis masekelai Souza-Gonçalves et Lopes-Andrade

37’ Pronotum with anterior corners rounded……………………..………………………………………38

38 (37’) Scutellar shield about 2× as long as elytral megapunctures. Eastern NW, eastern and western KZN…………………....................................………….…Cis stalsi Souza-Gonçalves et Lopes-Andrade

38’ Scutellar shield about 4× as long as elytral megapunctures. EC, GP, KZN, LP, MP, NW and WC…………………………...….…...………………Cis neserorum Souza-Gonçalves et Lopes-Andrade

39 (28’) TL usually less than 1.80 mm; IF about 1.80 mm, THEN elytral vestiture consisting of short bristles (0.01–0.02 mm)...………………………………………………...…………………………………..40

39’ TL more than 1.80 mm AND elytral vestiture consisting of moderately short to moderately long bristles (at least 0.04 mm)………..……………………….…………………...……………………………..43

40 (39) Elytral vestiture seriate, consisting of moderately long setae (0.04–0.05 mm). Southern WC…………………………………………………………………….Cis regius Orsetti et Lopes-Andrade

40’ Elytral vestiture non-seriate, consisting of short bristles or minute setae (≥0.02 mm)………...…..41

41 (40’) Pronotum devoid of impunctate median line; punctures separated by two puncture-widths or less; lateral margins crenulate. Southeastern MP, northeastern NW and western KZN………………………………...……Cis foveocephalus Souza-Gonçalves et Lopes-Andrade sp. n.

41’ Pronotum bearing impunctate median line; punctures usually separated by one puncture -width or less; lateral margins not crenulate………………………………...……………………………………..42

42 (41’) Dorsal vestiture consisting of thin and sparse setae. Southern WC, southeastern and western KZN……...................................…..……….Cis parvisetosus Souza-Gonçalves et Lopes-Andrade sp. n.

42’ Dorsal vestiture consisting of thick and dense bristles. Eastern and southeastern MP, northern and southeastern LP……..………...Cis mpumalangaensis Souza-Gonçalves et Lopes-Andrade sp. n.

43 (39’) Pronotum with punctures separated by less than half a puncture-width; lateral margins barely to completely visible from above. Elytra with single punctation. Cosmopolitan; introduced at southern WC..…………..…..………….……..…………..…..…………..…..…..…Cis chinensis Lawrence

43’ Pronotum with punctures separated by one to two puncture-widths; lateral margins not visible from above. Elytra with inconspicuously dual punctation. Eastern and southeastern KZN….….………………………..…………………Cis paraliacus Souza-Gonçalves et Lopes-Andrade

4. Discussion

The continental and insular sub-Saharan Ciidae faunas are more diverse than the northern fauna. The northern African fauna encompass 14 described species in four genera: Cis (10 species), Hadreule Thomson (1), Orthocis Casey (1) and Ropalodontus Mellié (2); and most of these species, if not all, are shared with the European fauna. The continental and insular sub-Saharan faunas encompass 86 described species in eight genera: Ceracis Mellié (1 introduced species), Cis (63, of which 11 were added in the present work), Dimerapterocis Scott (1), Ennearthron Mellié (1), Orthocis (6), Paratrichapus Scott (1), Tropicis Scott (4) and Xylographus Mellié (9).

Cis is the most diverse genus in southern Africa (comprising Botswana, Lesotho, Namibia, Republic of South Africa, Swaziland and the southern tip of Mozambique), and the great majority of species are known to occur in one or few provinces of Republic of South Africa. Cis neserorum is the unique exception occurring in seven of nine South African provinces [11]. Besides C. neserorum that is distributed from the south throughout the northeast of Republic of South Africa, the other species are more distributed in the northeast, with just eigth species occurring only in the southeast (C. afer, C, caffer, C. capensis, C. chinensis, C. muriceus, C. pickeri, C. regius and C. westerncapensis) and two occurring in the south and northeast (C. fuscipes and C. parvisetosus) (Figure 13, Figure 14 and Figure 15) (see distributional maps in Souza-Gonçalves & Lopes-Andrade [11,12,13]). It is important to note that most southern African Cis are restricted to these areas.

The southern African ciid with the largest distribution is the invasive species Cer. tabellifer [48]. The distributional patterns of southern African Cis species could provide insights about the possible impacts of the invasive species Cer. tabellifer in the autochthonous fauna, a hypothesis under evaluation by us and to be treated in a forthcoming paper. However, we can already highlight that there is likely a displacement of autochthonous Cis species to the northeastern of the Republic of South Africa (Figure 13, Figure 14 and Figure 15) (see distributional maps in Souza-Gonçalves & Lopes-Andrade [11,12,13]).

Except for the South African fauna, of which knowledge has been increased in the last years [1,10,11,12,48], information about the southern African fauna as a whole is very poor. The available information for the rest of this area is restricted to the original descriptions only and designations of lectotypes [21,22,23,24,25,49], without any further records in the scientific literature. Recent field collections and examination of historical collections have resulted in new discoveries about the African fauna, as undescribed genera (one from Tanzania and another from Madagascar), and records of undescribed species of Notapterocis Lawrence and Aliocis Sandoval-Gómez & Lopes-Andrade from the Republic of South Africa, which are being described by us.

Some species have been redescribed [27] and new species have been found in historical collections [13]; however, a gap in the literature remains concerning species from Central Africa and some from southern Africa. In most described species, there is no information on the morphology of sclerites of male and female abdominal terminalia, and specially the morphology of male genitalia, which is crucial for defining species boundaries and identifying them.

5. Conclusions

The southern African Cis fauna, comprising Botswana, Lesotho, Namibia, Republic of South Africa, Swaziland and the southern tip of Mozambique, now encompass 29 species: C. afer, C. aster, C. bicaesariatus, C. bimucronatus, C. caffer, C. capensis, C. chinensis, C. delagoensis, C. foveocephalus, C. fuscipes, C. grobbelaarae, C. lacinipennis, C. makebae, C. makrosoma, C. mandelai, C. masekelai, C. mooihoekite, C. mpumalangaensis, C. muriceus, C. neserorum, C. paraliacus, C. parvisetosus, C. regius, C. stalsi, C. tessariplacus, C. testaceus, C. urbanae, C. umlalaziensis and C. westerncapensis. We conclude that examination of sclerites of male terminalia is the most accurate way for defining species boundaries and identifying them, mainly those of the neserorum and pacificus groups, of which the species are externally very similar to each other. The knowledge concerning the few Central African and southern Africa species described in the XIX and beginning of the XX centuries is very poor, and it is still necessary to examine type material and provide information and images, especially of their male genitalia.