A new genus from Vietnam (Hymenoptera, Braconidae, Opiinae), and the description of two new species

Two aberrant species of Opiinae are described and illustrated from Vietnam, of which one is included into a new genus (Canalopius gen. n.; type species: C. periscopus sp. n.). The other species, Xynobius chrysops sp. n. belongs to an aberrant group of species near X. maculipennis (Enderlein, 1912).

Xynobius is a fairly large genus, of which many described species remain wrongly classified in Opius. Another problem is the undercollecting of this group; we have seen many new species from the Palaearctic and Oriental regions in the few recently made collections. One of these collections is from Vietnam assembled by a joint effort of Naturalis Biodiversity Center (RMNH, Leiden) and the Institute of Ecology & Biological Resources (IEBR, Hanoi). During a short visit to Leiden the first author sorted the two highly aberrant species of Opiinae described in this paper.
Opiinae are solitary koinobiont endoparasitoids of dipteran larvae. The parasitoid larva has its final development when the host larva has made its puparium; after pupation the adult wasp emerges from this puparium (Shaw and Huddleston 1991).

Material and methods
The material examined is deposited in the collection of Naturalis Biodiversity Center (RMNH) at Leiden. The specimens were collected in alcohol 70% using Malaise traps and the specimens were later prepared with the AXA method (van Achterberg 2009) and card-pointed.
For identification of the subfamily Opiinae, see van Achterberg (1990Achterberg ( , 1993, for identification of the genera, see Wharton (1997Wharton ( , 2009, Chen and Weng (2005), Li et al. (2013), Tan et al. (2016) and the diagnosis of the new genus in this paper. For references to the biology, see Yu et al. (2016) and for the morphological terminology used in this paper, see van Achterberg (1988van Achterberg ( , 1993, including the abbreviations for the wing venation. Measurements are taken as indicated by van Achterberg (1988): for the length and the width of a body part the maximum length and width is taken, unless otherwise indicated. The length of the mesosoma is measured from the anterior border of the mesoscutum till the apex of the propodeum and of the first tergite from the posterior border of the adductor till the medio-posterior margin of the tergite.
Descriptions and measurements were made under a stereomicroscope (Zeiss Stemi SV 6). Photographs were made with a Keyence VHX-2000 digital microscope.
Metasoma. Length of first tergite 1.1 times its apical width, convex and irregularly rugose medio-posteriorly and remainder largely smooth (Fig. 6), dorsal carinae strong basally and reaching apex of tergite, dorsope absent; second suture almost invisible; basal depressions of second tergite shallow, wide and oblique, and second tergite 0.8 times as long as third tergite; second and following tergites smooth, shiny and with row of setae posteriorly; combined length of second and third metasomal tergites 0.3 times total length of metasoma; setose part of ovipositor sheath 0.1 times as long as fore wing (entire visible sheath 0.14 times), 0.5 times first tergite (entire sheath 0.9 times) and 0.3 times hind tibia (entire sheath 0.4 times); hypopygium nearly 0.2 times as long as metasoma, truncate apically and not reaching apex of metasoma (Fig. 11).
Head. Antenna with 35 segments, bristly setose and 1.3 times as long as fore wing; third segment 1.1 times as long as fourth segment, length of third, fourth and penultimate segments 5.0, 4.5 and 2.3 times their width, respectively (Figs 12,22); length of maxillary palp 1.2 times height of head; length of eye in dorsal view as long as temple (Fig. 21); temple in dorsal view shiny, smooth, temple and vertex with adpressed setae; OOL: diameter of ocellus: POL= 1:1:3; face sparsely punctate, with a medio-longitudinal ridge extending to level of antennal sockets (Fig. 20); frons glabrous behind antennal sockets; in front of anterior ocellus shiny, smooth and glabrous but laterally setose (Fig. 20); labrum invisible; clypeus nearly semi-circular, with some oblique striae, convex but flattened ventrally, and its ventral margin truncate and narrow (Fig. 20); clypeus 2.1 times wider than its maximum height and 0.6 times wider than face; hypoclypeal depression narrow, slit-shaped; mandible straight ventrally, hardly twisted, apically gradually narrowed and second tooth small; mandible and with narrow ventral carina (Fig. 20); occipital carina remains far removed from hypostomal carina ventrally and horizontal dorsally, narrowly interrupted medio-dorsally; hypostomal carina narrow; malar suture distinct, narrow; length of malar space 0.9 times basal width of mandible.
Metasoma. Length of first tergite 1.7 times to its apical width, convex, its surface irregularly rugose medially (Fig. 19), dorsal carinae strong and combined in its basal third and area below widely depressed, but dorsope small; second suture almost invisible; basal depressions of second tergite minute and tergite as long as third tergite, both with wide setose bands (Fig. 19); second and following tergites smooth, shiny and setose posteriorly, especially densely in third tergite; combined length of second and third metasomal tergites 0.4 times total length of metasoma; setose part of ovipositor sheath 0.07 times as long as fore wing (exposed sheath 0.11 times), 0.6 times first tergite (entire sheath as long as tergite), and 0.2 times hind tibia (entire sheath 0.4 times); hypopygium about 0.3 times as long as metasoma, truncate apically and not reaching apex of metasoma (Fig. 23).

Biology. Unknown.
Notes. This aberrantly coloured new species belongs to a small group of Asian spp. near Xynobius maculipennis (Enderlein, 1912) united by the subbasally widened hind tibia with the resulting small knob glabrous and shiny dorsally, the hypopygium dark brown, the head and mesoscutum densely pubescent, and the fore wing with a large Y-shaped dark brown area below para-and pterostigma (Fig. 15). The new species differs from all other species examined by having the third metasomal tergite with a wide band of setae apically ( Fig. 19; narrow in other species); fore wing infuscate apically ( Fig. 15; subhyaline); the hypoclypeal depression nearly absent (distinct); the head and mesosoma dark golden pubescent combined with a slender first tergite (slightly widened apically and 1.6 × longer than its apical width; other species have usually pale yellowish or silvery pubescence, if dark golden than first tergite 1.2-1.3 times as long as wide apically).

Introduction
Pompilidae is a megadiverse family of ectoparasitoid wasps, with more than 5,000 valid species distributed worldwide and approximately 1,000 species inhabiting the Neotropical region (Shimizu 1994, Pitts et al. 2006. Although some pompilid species show primitive levels of sociality, most species are solitary (Shimizu 2004, Shimizu et al. 2010. These species are known as "spider wasps", since adult females usually uses spiders as food resource for their larva (Brothers andFinnamore 1993, Wasbauer 1995). Pompilids presents a wide diversity of behavioural strategies to parasitize their hosts, with species nesting in the ground, in pre-existing cavities, mud nests they have built, or using as hosts active spiders that are posteriorly killed by the wasp's larvae (El-Hennawy 1996, Souza et al. 2015. Most species are solitary, but some show primitive levels of sociality (Shimizu 2004, Shimizu et al. 2010).
The genus Epipompilus Kohl comprises 52 species of spider wasps, 16 of which occur in the Neotropical region, one in the Nearctic region, and 36 species in the Australian region (Roig-Alsina and Barneche 2017). The current biological knowledge of this genus is based on three species from the Australian region, E. insularis Kohl, 1884, E. exleyae Evans, E. reiki Evans (Evans 1972, Pollard 1982, Harris 1987, and one Neotropical species, E. platensis (Roig-Alsina and Barneche 2017). Evans (1972) reported E. exleyae as an ectoparasitoid of immature Sparassidae, and E. rieki as an ectoparasitoid of an unidentified spider. Pollard (1982) and Harris (1987) reported E. insularis as a koinobiont ectoparasitoid of multiple cursorial spiders, including the clubionid Clubiona cambridgei Koch, 1873, and the salticids Salticus sp., Trite auricoma Urquhart, 1885 and T. planiceps Simon, 1899.
Segestriidae (Simon, 1893) consists of haplogyne spiders, small-to-medium in size (total body length between 2.8 and 20.25 mm) (Grismado 2008, Giroti andBrescovit 2018), usually found in loose barks, cracks in tree trunks, under rocks, or even in anthropomorphized places (Roig-Alsina and Barneche 2017). These spiders are nocturnal, sedentary, construct funnel-like webs and have been reported as a host of Epipompilus platensis (Roig-Alsina and Barneche 2017). Currently, Segestriidae comprises 130 species divided into four genera (World Spider Catalog 2018), among them Segestria Latreille, 1804 and Ariadna Audouin, 1826 occurs in South America (Grismado 2008, Giroti and Brescovit 2011, World Spider Catalog 2018. The genus Ariadna currently consists of 105 of the 130 species included in Segestriidae (Giroti and Brescovit 2018). This genus is distributed across all continents, but the greatest number of species is found in tropical and subtropical regions. The knowledge of biology and ecology of Ariadna is quite incipient, based on a single study that reported A. maxima as host of the koinobiont ectoparasitoid Sphaerops appendiculata Philippi, 1865 (Diptera, Acroceridae). This acrocerid is the only external koinobiont parasitoid of spiders known in the family, since other species develops inside the opistosoma of spiders (Schlinger 1987).
Here, we report E. excelsus (Bradley, 1944) as a koinobiont ectoparasitoid of the tube-dwelling spider Ariadna mollis (Holmberg, 1876), with behavioural and biological notes of this wasp-spider interaction.

Methods
We observed E. excelsus attacking A. mollis on May 5, 2017 at Serra do Japi (23°13'53.60"S, 46°52'47.01"W), a tropical forest located in the municipally of Jundiaí, São Paulo state, Brazil. After the wasp attack, we collected the parasitized spider and transported it to the Laboratório Especial de Coleções Zoológicas (LECZ) -Instituto Butantan, São Paulo, Brazil, where we recorded the larval development. We kept the spider inside a terrarium (15×15×13 cm) and provided cotton balls with water, a piece of loose bark from the tree that we found the spider (used as a shelter by the spider), and we fed the spider every two days with Chrysomya putoria (Wiedemann) (Diptera: Calliphoridae). To access the wasp development (i.e. from the egg stage to adult emergence), we photographed the parasitized spider daily upon a mm scale. We used the software Adobe Photoshop CS6 to analyze the photographs and measure the length and width of the larvae throughout its development. We deposited the remains of the spider carcass and the adult parasitoid in the Arachnida collection of Instituto Butantan, São Paulo, Brazil (Curator: A. D. Brescovit).

Results and discussion
We found an adult female of E. excelsus in an inspection flight near a trunk of a Plinia cauliflora Mart. (Myrtaceae). The wasp found an adult female of A. mollis and flew around the spider for few (six minutes approximately) minutes, remaining at a distance of approximately 6 cm from the spider. The wasp then landed on the spider dorsum, stung and paralyzed the spider. Thereafter, the wasp laid an egg dorsally on the spider abdomen; oviposition took approximately three minutes. Then the wasp drummed the spider (Fig. 1A), and after five minutes, flew off leaving the parasitized spider paralyzed. After approximately two more minutes, the individual of A. mollis began to move slowly and continued its normal course on the tree trunk (Fig. 1B).
In laboratory, the parasitized spider constructed a tube-shaped shelter in the space between the terrarium wall and the piece of bark. The spider continued to feed on the flies normally during our observations. The egg developed in two days. Once it hatched, the first instar larva emerged and stuck to the spider abdomen. The larval development included four instars distinguished by abrupt increases of the larvae size between each larval stage. The first instar measured 0.3 mm in width and 2.2 mm in length (Fig. 1C). The larvae reached the second instar (1.1 mm wide and 4.1 mm long,   1D) five days later. After three days, the larvae reached the third instar and measured 1.7 mm wide and 6.1 mm long (Fig. 1E). Finally, the fourth instar was observed five days later, measuring 2.2 mm wide and 7.8 mm long, and presented 11 retractile dorsal tubercles (Fig. 1F). The fourth instar larvae killed the host and slowly consumed more than 50% of its body. Only the legs, the part of the ventral abdomen containing the genitalia, and the anterior part of the cephalothorax were not consumed by the larvae (Fig. 1G). On the next day, the larva began to construct a cocoon in the space between the bark and the wall of the terrarium, and fixed the apical portion of the cocoon in the bark (Fig. 1H). Emergence of the adult wasp occurred by the apical region of the cocoon five days after cocoon construction. The entire developmental cycle from oviposition until the emergence of adult wasp, took 24 days. This period is shorter than the developmental periods of wasps of other Pompilidae species. Paracyphononyx scapulatus, for example, develops in 54 days (Souza et al. 2015), P. rufricus in 30 days (El-Hennawy 1996), and Notocyphus tyrannicus Smith in 46 days (Martins et al. 2016).
Studies on congeneric species do not present a detailed description of the individual's life cycle (Roig-Alsina and Barneche 2017). Studies on the interaction between the parasitoid wasp Epipompilus insularis and the spiders Clubiona cambridgei, Trite auricoma or T. planiceps (Harris 1987) and the interaction between E. insularis and the spiders C. cambridgei or Salticus sp. (Pollard 1982) showed similar results to those of the present study. Harris (1987) and Pollard (1982) suggested that this genus of diurnal parasitoid wasps usually parasitize nocturnal non-weaver spiders because these spiders rest in shelters during the day, since this behavior of inactivity during daytime decreases the risk of attacks to the spiders, and thereby provides protection for the wasp egg and larvae. It is not possible to predict whether the parasitized spiders return to its shelters, but notably, our study specimen built a shelter and returned to it every day after goes out at night to capture the preys we offered. The spider was active during almost the entire larval development of E. excelsus, sheltering only in the final phase of development (fourth instar), before the wasp kill and consume the host. Harris (1987) reported a similar life cycle development time for E. insularis (22 days total) when compared to E. excelsus (24 days). The time of development time of each larval instar was also similar. However, the larval development of E. insularis involves five instars, while our study species presented four instars. Moreover, the final larval instar of E. insularis presents 12 lateral tubercles, while E. excelsus has 11 tubercles at the last instar.
Although all Pompilidae species use spiders as hosts, there are a high diversity of behavioural strategies characterized by different ethological sequences according to Evans (1953). This author classified the behaviour of E. insularis as VPO (Venari = to hunt; Pungere = to sting the prey; Ovum parere = to lay the egg) or VO (Venari= to hunt; Ovum parere= to lay the egg), and discussed that such ethological sequences represent primitive conditions within Aculeata, but derived conditions within Pompilidae. Species of other genera, such as Notocyphus Smith, 1855, Paracyphononyx Gribodo, 1884 and Homonotus Dahlbom, 1843 present the same ethological sequence (Evans 1953, Souza et al. 2015. In conclusion, this study showed that the Neotropical species E. excelsus presents similar biological characteristics (e.g. koinobiont ectoparasitoidism) when compared to the Australian species. On the other hand, there are differences in larval morphology (e.g. number of tubercles) and development (e.g. number of larval instars). Studies on other koinobiont ectoparasitoid species would elucidate the extent of these differences and similarities, contributing to our understanding of the evolutionary history of Epipompilus, and consequently of Pompilidae.

Introduction
Pollinators are a key component for global biodiversity, because they assist in the sexual reproduction of many plant species and play a crucial role in maintaining terrestrial ecosystems and food security for human beings ( Tropical dry forest present a great bee species richness in Mexico (Ayala 1988, Ayala et al. 1996, which is explained by the high diversity of entomophilous plants (Ayala 2004), and their differentiated phenology (generally, tree species flowering in dry season while herbaceous and shrub species flowering in wet season), which provides to this group of organisms, resources for most of the year (Reyes-Novelo et al. 2009). This vegetation type covered approximately 270,000 km 2 in 1990 but currently there are great pressures exerted on this type of forest. For example Trejo and Dirzo (2000), reported that 50% was either fragmented or existed as relicts. Furthermore, they found changes in land use on 23% of the surface and that only 27% of this type of vegetation remained intact (Trejo and Dirzo 2000).
Land use change, such as degradation or fragmentation of tropical dry forest, could have consequences on the richness and composition of bee communities. According to previous studies, bees are susceptible to the loss of their natural habitats by the fragmentation and transformation of the landscape for agricultural, livestock and urban purposes (Martins et al. 2013, Woodcock et al. 2013, Le Féon et al. 2016, Mogren et al. 2016. For this reason, it is important to compare local bee communities from disturbed environments with one from a preserved environment in the same region, as this should provide useful information for making conservation-based decisions on bee community management (Dalmazzo 2010).
Two of the most important factors for bee communities are food availability (nectar and pollen) and nesting sites (Reyes-Novelo et al. 2009). Changes in land use are expected to affect both and consequently alter the structure of these communities. Due to this, the aims of this study were: 1) to ascertain whether landscape transformations affect bee community structure (richness, abundance, and α and β diversity) and, 2) to identify which are the bee groups most susceptible to these changes, as well as those who benefit from them.

Study area
The study is located at Área de protección de flora y fauna Sierra de Quila, (here and after APFFSQ) and its zone of influence, which is found in the south-central portion of Jalisco, Mexico ( Fig. 1). It is located within the Trans-Mexican Volcanic Belt physiographic region. The research was conducted in the municipalities of Tecolotlán, Tenamaxtlán, San Martín Hidalgo and Cocula. Geographically, it is found between 20°14' and 20°22'N; 103°57' and 104°07'W (DMS), with an area of 15,912 hectares (Villavicencio et al. 2012).

Data collection
Three conditions in tropical dry forest (TDF) with different land-use were selected. The first was an urbanized area (U) at Tecolotlán, a municipality with 9,189 inhabitants (IIEG 2015); the second was an agricultural area with crops and livestock (CL), which begins on the area surrounding the town, and is characterized by the presence of crops (mainly corn -Zea maysm L.) as well as cattle ranches, which were slightly fragmented by secondary vegetation and some remaining trees of the TDF, used as living fences. The third was an area of preserved vegetation (P) of the TDF within the APFFSQ. Some of the most common trees in this area are from the following genera: Bursera, Ceiba, Erythrina, Heliocarpus, Lysiloma, Stenocereus and Viguiera ( Fig. 1) (Guerrero-Nuño and López-Coronado 1997).
Sampling of wild bees was conducted throughout the year, with three sampling days per month (one working day per condition, from 10:00 to 16:00 hours). Four plots, 50 × 5 m, were randomly located for each land use condition and the bees were sample walking along the transect during periods of 60 minutes per collection. Bee species presences and their abundance were recorded. Bee species that could not be determined in the field were collected using an entomological net with extension of 2.70 m on tree strata and an extension of 1.00 m for herbaceous strata. Specimens were processed according to Michener et al. (1994). Bees were then identified as precisely as possible with the help of the specialized literature, whether at a species or genus level, and then separated by morphospecies. The specimens were deposited at the Entomological Collection of Centro de Estudios en Zoología of Universidad de Guadalajara (CZUG).

Data analysis
Sampling efficiency and richness comparison A species accumulation curve was performed to evaluate the sampling effort per site and, as well as to compare the richness among the different conditions using ESTIMATES 9.1.0. (Colwell 2016). A rarefaction curve was also prepared using the same program.

Bees abundance
Bee abundances with and without Apis mellifera were compared via analysis of variance (ANOVA) for the first case, and Kruskal-Wallis followed by the Nemenyi post-hoc test, for the second case. Furthermore, Pearson's chi-squared test and residual analysis were used to compare the abundance of different bee families among the land-use conditions using the R 3.2.5 Sofware (R Core Team 2016).

Diversity analysis
The Shannon-Wiener index was used to compare alpha diversity of sites via a confidence interval obtained using bootstrap in PAST 3.15 software (Hammer et al. 2001). In order to contrast the composition of tribes in the study areas, beta diversity index (ßsim) was estimated, while non-metric multidimensional scaling (NMDS) was used with the R 3.2.5 software (R Core Team 2016) via the 'vegan' package (Oksanen et al. 2013).
The richest condition was P, with 120 species, followed by CL with 98 species and, finally U, with 81 species. The rarefaction curve among different land-use conditions indicated greater richness in P than in U and CL, although they did not present statistically significant differences (Fig. 2). Furthermore, a sampling efficiency of 88% was obtained for P (135 spp.), 77% for CL (127 spp.), and 79% for U (102 spp.) in accordance with the richness estimated by Chao 1).

Abundance of bees
The highest number of bee individuals was observed in P (5,067), followed by CL (4,786) and U (4,202). The ANOVA (df = 2, F = 0.352, p = 0.704) showed no significant differences between the registered abundance of bees for the different types of land-use. If only wild bees are considered (excluding Apis mellifera), the greatest abundance was in P (2,731), followed by U (2,418) and, finally, CL (1,876). In this case, Kruskal-Wallis test (df = 2, K = 7.265, p = 0.026) indicated significant differences between median abundances, while the Nemenyi post-hoc test demonstrated significant differences (p = 0.023) only in terms of the abundances of bees between P and CL.
Pearson's chi-squared test showed a dependency between land-use and bee family abundance (df = 8, χ 2 = 560.2715, p < 0.05) ( Table 2), while residual analysis indicated that abundance of Colletidae, Andrenidae and Megachilidae are positively related to P. The Halictidae was primarily associated with U where they were most abundant, while the Apidae registered the highest abundance in CL (Fig. 3).

Diversity analysis
The Shannon-Wiener index (H') values presented significant differences with a p < 0.05, P obtained a greater level of diversity compared to the other types of land-use, followed by U and, finally CL. Evenness (J') is lower in CL, while U and P presented similar even-   ness. Moreover, when only wild bees were considered, diversity was higher in P. This is due to the elevated abundance of Apis mellifera ( Table 3). The beta diversity index (βsim) showed an average species turnover rate of about 32% for each condition. In terms of bee species composition change, the highest was between U and CL (36%), followed by P and U with 30% and finally between CL and P with 27%.
The indirect ordination performed with NMDS resulted in a stress value of 0.1326, that shows a gradual separation of the three different land-use conditions along the first axis, where on the left side are located U sites, CL sites in the center, and P sites on the right side. The tribe Anthophorini and tribes of the Halictidae were associated with the U, while tribes of the Apidae and one megachilid tribe were located principally between the U and CL areas. The Lithurgini, two tribes of the Andrenidae and two cleptoparasite tribes (Ericrocidini, Epeolini) were associated with the CL, while the Xylocopini was associated with the P and CL areas. Tribes of Colletidae, Andrenidae, and Megachilidae and two apid tribes were found in P (Fig. 4).

Discussion
The highest bee richness was registered in P while the only significant differences were recorded between U and P. The results indicate a gradient in richness related to the habitat conservation. This finding is supported by different studies which show that the destruction of the habitat, for agriculture or urbanization, is principally responsible for bee species reduction, up to the point of local extinction for some species (Frankie et al. 1997, Martins et al. 2013. Furthermore, in a meta-analysis, Winfree   (2008), found that on average the richness of pollinators (principally bees) in crops decreased by half as the distance from preserved habitat increased to 1500 m. The decrease in bee richness is related to their dependence on various resources to complete their lifecycle, such as food resources (pollen and nectar) and nesting substrates (either from the soil or vegetation), and materials for constructing their nests (Winfree 2010). The reason that a greater number of bee species is registered in the preserved area is possibly due to a greater availability of plant species for food, being that bee richness has been positively related to the richness of flowers (Carper et al. 2014) and to a great floral diversity, which provide food to a great variety of specialist and generalist bee species (Martins et al. 2013).
The abundance of bees did not differ significantly between the three land-use conditions. This agrees with various studies that report a similar or greater abundance in human modified areas (Carper et al. 2014, Mallinger et al. 2016, Nantes-Parra et al. 2008, Olalde-Estrada et al. 2015, Otero and Sandino 2003, Tonhasca et al. 2002; this occurs in sites with moderate disturbance with gaps on forest canopy, which facilitate grow of herbaceous layer (Carper et al. 2014). This also shows that urban areas could be important for bee communities, despite the modification of vegetation, because they offer ornamental and wild flowers, particularly because they are watering them in times when native vegetation is diminished, as well as nesting resources associated to human constructions (Dalmazzo 2010). The differences in abundance between the P and CL when Apis mellifera is excluded from analysis could be due to the same reasons that affect bee richness in addition to the use of pesticides in the crops which affect wild bees (Winfree 2010).
The number of individuals per family varied, Andrenidae, Megachilidae and Colletidae being far less abundant than the Apidae and Halictidae. Potts et al. (2003) found the Andrenidae positively associated with annual plant diversity, and Megachile with plant richness, pollen and available nectar. Vergara (2005) suggests that, in areas with agricultural activity, bee genera that nest in the soil, including andrenids and colletids, are under-represented due to plowing for agricultural activities and the presence of livestock, which affect nesting. While Halictidae presented greater abundance in the urban area, this is possibly because most of the registered bees of this family are small (<1 cm) and most of them are polilectic (Michener 2000).
Banaszak-Cibicka and Zmihorski (2012), reported that urban areas contain small green isolated island (parks and gardens), that promote the abundances of small bees, which require only a small amount of food. Winfree et al. (2009), indicate that they do not have problems in finding nesting sites in disturbed areas.
Total bee abundance tends to decline at disturbed sites, while individual species present different responses, some of them show drastic reduction in their populations, while others remain stable or even increase with land-use change (Winfree et al. 2008). For this reason, the use of the habitat according to bee species plays an important role. Bees that require specific conditions for their development, become the most vulnerable to the disappearance of their habitat. On the other hand, if the resources generated by the modification of the habitat is exploited by the bees, they could survive in disturbed areas (Winfree 2010). That is the reason why bees that can use a wide variety of resources are often able to compensate when circumstances change, although some species disappear due to land use changes. The higher richness and evenness patterns in preserved areas is supported by Reyes-Novelo et al. (2009), who explain that in disturbed areas some species disappear locally and the abundance of others decreases, which modifies dominance.
Finally, β diversity showed a medium level of species turnover rate among conditions, despite the maximum distance between sampling sites was 6 km. The structure of bee community contrasts with P and U, in which tribes such as Halictini and Anthophorini (due to the high abundance of Anthophora squammulosa) are associated with the urban area. The tribes of the Andrenidae, Colletidae, Megachilidae and Apidae, such as bumblebees (Bombini), are strongly associated with P, while the CL area is found at the mid-point of the gradient. This suggests that disturbance, with its variations in micro-environments and food and nesting resource has a significant effect on bee community species composition.
The bee fauna of the TDF in Mexico is one of the richest and most diverse in the country (Ayala 2004), so, the protection of this type of vegetation is crucial for the conservation of this group of insects, given that the reduction of these native pollinators could have a negative economic impact on crops (Maass et al. 2005). As the expansion of human settlement and the population growth or production activities have led to the process of deforestation, fragmentation and degradation of this forest (Maass et al. 2010), Trejo andDirzo (2000) report that approximately 3.7% of the TDF of the total area of Mexico (72,850 km 2 ) remains relatively intact and with a high annual deforestation rate of 2.02% of the total coverage.

Conclusions
The APFFSQ is an important site for maintaining wild bee diversity, given that 200 species of bees were registered in the area, of which 160 are present in the TDF (Razo-León 2015). According to Ortega, (2007), the TDF covered 19.84% of APFFSQ (2,797 ha) in 1993, by the year 2000, it only covered 15.25% (2,149 ha), this means that the TDF lost 23.2% of its area in seven years, because it is rounded by agricultural or livestock production areas which make it more vulnerable to transformation. This leads to the conclusion that change in land-use negatively affects the richness, abundance, and diversity of species of bee community, for this reason it is recommended that the TDF in Mexico should be protected in order to guarantee the conservation of this group of insects.  (Michener 2007). The present paper is part of a series of works dealing with the bees of the genus Sphecodes of the territory of the Palaearctic region , 2015a, b, c, 2016a, b, 2017a, b, c, 2018, 2018a. Currently 77 species are known from this region, but the Sphecodes fauna of Iran is particularly under-recorded.
There are currently 862 species of bee known from Iran (Ascher and Pickering 2018), but new records and species continue to be added (e.g., Khodaparast and Monfared 2012, Khaghaninia et al. 2013, Nadimi et al. 2014Kuhlmann and Proshchalykin 2015, Safi et al. 2018, Proshchalykin and Kuhlmann 2018. The genus Sphecodes has been one of the more overlooked taxa partly due to the complexity of positively identifying material. Only one species -Sphecodes persicus Blüthgen, 1924(=S. pinguiculus Pérez, 1903 of this genus has been described from Iran (Blüthgen 1924) so far, and in total only 16 species have been recorded for the country (Warncke 1992, Ascher and. Records of Sphecodes from Iran in other published sources Straka 2012, Özbek et al. 2015, Astafurova and Proshchalykin 2017b) were not original and referenced from a previous paper (Warncke 1992).
Here we have attempted to build a survey of Iranian material available and to provide an overview of the 25 species found across Iran. Of these species, eight are recorded for the first time for the Iranian fauna and one species is described as new. For each species we provide a list of Iranian localities from the material examined and give previously published occurrences along with a summary of the wider distribution.

Materials and methods
The results presented in this paper are based on 324 specimens collected in Iran and currently housed in the Zoological Institute (St. Petersburg, Russia) and private collection of Maximilian Schwarz (Ansfelden, Austria). Acronyms for collections from which specimens were borrowed or are deposited are as follows: OÖLM -Oberösterreichisches Landesmuseum, Linz, Austria; PCMS -private collection of Maximilian Schwarz, Ansfelden, Austria; ZISP -Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia.
The taxonomy and distribution of species generally follows that of Warncke (1992), Bogusch and Straka (2012), Astafurova and Proshchalykin (2017b), and Özbek et al. (2015). A detailed current synonymy of the species has been given by Astafurova and Proshchalykin (2017b). Morphological terminology employed below follows that of Michener (2007) and Engel (2001). The ventral surfaces of some flagellomeres bear a distinctive patch or zone of sensilla trichodea A (sensu Årgent and Svensson 1982), and we refer to these as the "tyloids", easily observable under light microscopy. The abbreviations F, T, and S are used for flagellomere, metasomal tergum, and metasomal sternum, respectively. The density of integumental punctures is described using the following formula: puncture diameters (in μm) / ratio of distance between punctures to average puncture diameter, e.g., 15-20 μm / 0.5-1.5. Integumental sculpturing, aside from distinctive surface punctation, is described as follows: reticulate: superficially netlike or made up of a network of raised lines; rugose: irregular, nonparallel, wrinkled raised lines (rugae); tessellate: a regular network of shallow grooves with flat interspaces.
Specimens were studied with the use of a Leica M205A stereomicroscope and photographs taken with a combination of stereomicroscope Olympus SZX10 and digital camera Canon EOS70D. Final images representing a composite of several photographs taken at different focal planes and combined using Helicon Focus 6. All images were post-processed for contrast and brightness using Adobe Photoshop.
The species are presented alphabetically and those that could not be inspected in this paper are quoted from published sources. Provinces are presented in alphabetical order and the names of provinces are given in bold type. New distributional records are noted with an asterisk (*). Variation. Specimens from Iran differ from specimens from the rest of the range by having an entirely dark metasoma with distinctly punctate T2 in male. Diagnosis. The male is close to Sphecodes nomioidis Pesenko, 1979 owing to similar structure, sculpture of the body including distinctly punctate metasomal terga and similar small trapezoidal gonostylar shape (Fig. 1a, b). In gonostylar shape the new species is also close to S. geoffrellus (Kirby, 1802) ( Fig. 1 c), but differs from both species by having less developed tyloids covering about 1/2 ventral surface of flagellomeres (Fig. 1d) (covering at least 4/5 ventral surface of flagellomeres in S. nomioidis and S. geoffrellus) (Fig. 1e).

Sphecodes croaticus
Description. (Male). (Fig. 1f ). Total body length 5.5 mm. Head (Fig. 1g) black (except brown mouthparts and antenna); weakly transverse, at most 1.1 times as wide as long; vertex not elevated, distance from top of head to upper margin of a lateral ocellus about 2 lateral ocellar diameters as seen in dorsal view; antenna attain middle of mesoscutum, F1 transverse, 0.75 times as long as wide, F2 1.3 times as long as wide, remaining flagellomeres about 1.2 times as long as wide; tyloids (from F3 onwards) covering about 1/2 ventral surface of flagellomeres; clypeus, paraocular area and frons with confluent punctures (15-25 μm); area between ocellus and eye with punctures separated at most by 1.5 of a puncture diameter; vertex behind ocellus and gena rugose; face below antennal toruli with snow-white, plumose pubescence, obscuring the underlying integument.

Sphecodes rubicundus Hagens, 1875
S. tadschicus Blüthgen in Popov, 1935. Nevertheless records of this species in north-western provinces of Iran are certain in the future as it is known from Caucasus and Turkey. Distribution. Europe (north to Germany), Russia (European part), Caucasus, Turkey, Iran (?), Israel.

Discussion
Iran is a mostly mountainous country, with a landscape dominated by mountain ranges that separate various basins or plateaux from one another. A great variety of terrestrial ecosystems are situated on the territory of Iran: desert, semi-desert, steppe, forest steppe, forests and woodlands. However, dry biotopes occupy the majority of the Country and only 7% of the surface area is forested (most forest is found on the mountain slopes). Landscape and biotope diversity have a great effect on the diversity of bees, observable in the Sphecodes fauna of Iran, which includes forest species of temperate zoneas well as steppe and desert species of subtropics and endemic mountain species. The majority of the Sphecodes fauna of Iran is formed by 14 widespread species, distributed from Europe to West Siberia or even to the Russian Far East and Japan: These are S. albilabris, S. alternatus, S. crassus, S. ephippius, S. gibbus, S. longulus, S. monilicornis, in Central Asia and Iran. Sphecodes anatolicus is a rare mountain species known from Mediterranean and Caucasian Regions to Central Asia. Sphecodes olivieri is widespread in semi-desert and desert of the Western Palaearctic. Sphecodes ebmeri sp. n. is a mountain edemic of Elbrus which also could be found in mountain Turkey and the Caucasus. In total, 25 species of Sphecodes are recorded from Iran. This is distinctly less in comparison with the adjacent fauna of Turkey, Caucasus and Central Asia (Table 1). However this number will probably increase at least by a third and could be richer owing to eleven Western Palaearctic species which could be found in Iran: S. cristatus, S. hakkariensis, S. ferruginatus, S. nomioidis, S. dusmeti, S. niger, S. nurekensis, S. intermedius, S. armeniacus, S. crassanus, and S. geoffrellus. There are some species distributed in southern Europe, Turkey and the Caucasus that could be found in north-western Iran, whereas species mostly distributed in Central Asia could be recorded in central and eastern Iran. Because of these species the Iranian fauna of Sphecodes may be more diverse than those of Turkey or Central Asia.

Introduction
The genus Celonites is very distinct from other pollen wasps in the wide horizontal lamellae on the propodeum and the acute sides of the metasoma (Richards 1962). Both characters are related to the outstanding ability of the imagines to roll up (Gess and Gess 2010). Approximately 46 species of the genus (including six well separated subspecies) have been recognized in the Palaearctic region (according to the list of Carpenter 2001, combined with newly described or synonymized species by Gusenleitner 2002Gusenleitner , 2007Gusenleitner , 2012Gusenleitner , 2018Mauss 2013, Mauss et al. 2016. Within the Palaearctic Celonites abbreviatus-group (sensu Richards 1962) the Celonites abbreviatus-complex constitutes a well defined monophylum characterized by the existence of knobbed setae at the front of the head forming a pollen collecting apparatus (Mauss 2013). All members of this species-complex for which flower visits have been recorded were observed to visit nototribic flowers of Lamiaceae (Bequaert 1940, Schremmer 1959, Müller 1996, Mauss 2006, Mauss 2013, Mauss et al. 2016. During pollen uptake from these flowers a female would stand on the lower lip of the corolla, while she performs at a high frequency slight back and forth movements, rubbing her head over the nototribic anthers. In this manner pollen grains are removed from the anthers with the stiff knobbed setae on frons and clypeus of the female (Schremmer 1959, Müller 1996, Mauss 2006, Mauss et al. 2016). The species-complex includes six nominate species, namely Celonites abbreviatus (Villers), C. andreasmuelleri Mauss, C. hermon Gusenleitner, C. mayeti Richards, C. persicus Richards and C. tauricus Kostylev (= C. spinosus Gusenleitner) (Mauss 2013, Mauss et al. 2016).
Celonites hermon was described from a single female collected by Bytinski-Salz at Mount Hermon in the Golan Mountains by Gusenleitner in 2002. He separated the specimen from the "very similar" Celonites abbreviatus because of its extremely long proboscis that reaches the distal end of the abdomen (Gusenleitner 2002). As already pointed out by Mauss (2013) proboscis length can not be examined in most Celonites specimens, since the mouthparts are usually retracted, while the protruded proboscis of the type specimen is an exception. This may have misled Gusenleitner (2002) into believing, that the length of the proboscis is outstanding in C. hermon. In fact, the careful investigation by Schremmer (1961) had already demonstrated that the proboscis of C. abbreviatus clearly exceeds the tip of the metasoma. Moreover, Mauss (2013) referred to a female of C. abbreviatus in his collection with protruded mouthparts extending beyond the end of the metasoma as well. Therefore C. hermon does not distinctly differ from C. abbreviatus in the character that mainly substantiated its description and was considered to be a junior synonym of C. abbreviatus.
However, Mauss (2013) avoided formal synonymy of both taxa as the situation appeared to be more complicated. The club of the antenna of the type of C. hermon is completely orange, which is also the case in specimens of C. abbreviatus from the Balkan and Aegean populations whereas the antennae of Central and West European specimens of C. abbreviatus are much darker suggesting that the eastern populations might belong to a different subspecies or even species that could be identical with C. hermon. Subsequently Mauss et al. (2016) demonstrated that Celonites tauricus differs distinctly from C. abbreviatus in that the likewise orange antennal club of C. tauricus has always a dark tip, which is possibly related to different male signals during mating behaviour. Since the tip of the antennal club of the holotype of Celonites hermon is completely orange, it can not be identical with C. tauricus. The distribution area of Celonites abbreviatus extends from Northwest Africa over the Iberian Peninsula, Southern and Central Europe eastwards to the Balkan and the western Aegean. Celonites tauricus is allopatrically distributed with regard to Celonites abbreviatus and has been recorded from the Crimea, Kos, Rhodes, Cyprus and Asia Minor (Mauss et al. 2016, Mauss unpubl.) (Fig. 11), separating the locus typicus of C. hermon by more than 1000 km from the eastern margin of the distribution range of C. abbreviatus. Therefore C. hermon could be hypothesized to represent a relict population of C. abbreviatus in the Golan Mountains that was separated from the Balkan and Aegean populations of C. abbreviatus as a result of range expansion of C. tauricus. An alternative hypothesis is that Celonites hermon is an indigenous biospecies. In this case one would expect closer relationship to other members of the C. abbreviatus-complex from geographically adjacent areas in the Middle East, that is Celonites tauricus or Celonites andreasmuelleri.
Based on new material of Celonites hermon, the aim of this study is to reinvestigate the status of the taxon carefully, to characterize its taxonomic position within the genus Celonites and to describe the newly discovered male.

Material and methods
The specimens of Celonites studied belong to the public collection of the Upper Austrian State Museum (Linz, Austria) (OLML), as well as to the private collection of J. Gusenleitner (Linz, Austria) (JG). The specimens were investigated under a WILD M3 stereo microscope (maximum magnification 80 times). Measurements of the exoskeleton were made using an ocular micrometer (highest resolution 0.011 mm). Distances between the ocelli, the compound eyes and the mesonotum width were measured according to Eck (1978). The genitalia of all males were extracted after re-softening the specimens and were then studied in 80% ethanol or 85% glycerol. Nomenclature of male genitalia follows that of Birket-Smith (1981). Drawings were made with a drawing tube (WILD Type 308700). Micro-photos were taken with a Leica IC 80 HD camera mounted on a Leica MS 5 stereomicroscope. Multifocuspictures were generated with Leica Application Suite (LAS) software. The scapus of the antenna is referred to as antennal article A1 and the pedicellus as A2, the flagellum consists of the articles A3-A12. Gusenleitner, 2002 Figs 1-9 Material studied. Israel: Mt. Hermon, 33.416111N, 35.857500E, 1900m a.s.l., 17.07.1971, 1♀ (holotype, dbM No. 3579), leg. Bytinski-Salz, coll. JG. Syria: 40 km NE of Damascus, 33.416667N, 35.866667E, 13.05.1996West Syria, Tartus env., 34.900000N, 35.883333E, 25.05.1996, 2♀♀ (dbM No. 3691, 3692), leg. Mi. Halada, coll. OLML.

Celonites hermon
Diagnosis. Pale markings light yellowish to yellow without reddish tinges (Fig. 1). Immediate tip of antennal club (A8-12) and especially ventral side of A12 orange to orange-brown, not markedly contrasting in colour from adjacent light ventral side of antennal club. Axilla of mesoscutellum with short blunt lateral projection that only slightly projects over adjacent posterior part of tegula. Frons and clypeus covered with pale, stiff pollen collecting setae (Fig. 3). In females most, in males at least some of these setae with tiny spherical enlargement at tip ("knob"). Diameter of median ocellus distinctly larger than diameter of lateral ocelli, ocelli larger than in C. abbreviatus and C. tauricus. Median ocellus circular, lateral ocelli more or less obliquely oval (Figs 4, 7). Vertex falling away obliquely towards the occipital carina immediately behind ocelli (Figs 4, 7).
Females with head in front view more elongated in relation to its length. Clypeus with lateral margins more continuously rounded leading to elongated appearance (Fig. 2). Cuticula on frons and at least dorsal half of clypeus densely and strongly shagreened leading to a conspicuous dull region covered with knobbed pollen-collecting setae, changing gradually into the adjacent smooth, shiny cuticula on vertex and less densely shagreened cuticula ventrally on clypeus (Figs 2, 4). Knobbed pollen collecting setae on frons distinctly longer and relatively thinner in comparison with Celonites abbreviatus and C. tauricus, up to 1.4 times as long as diameter of median ocellus, with diameter almost equal along most of length, only weakly broadened towards base (Figs 3, 4).
Males with only two oval-shaped tyloids situated ventrally on articles A9 and A10 of club-shaped antennae. Midcoxa with a small but distinct spine at distal end, on anterior side of midcoxa close to its antero-medial angle. Tergum VII markedly lobed posteriorly (Fig. 6). Sternum VIII only shallowly emarginate, surface not much sunk in towards emargination (Fig. 8). Tergum VII with medial lobe distinctly emarginate leading to quadrilobed appearance of posterior margin (Fig. 6). Emargination of posterior margin of sternum VIII medially angled, laterally with a transverse section well set off by protruded edge with distinct little spine on each side of posterior margin (Fig. 8). Posterior margin of stipes of male genitalia with dorso-medial concavity (Figs 9, 10). Stipes in dorsal view elongated, with medial and lateral margin running more or less parallel towards posterior end (Figs 9, 10). Posterior process of volsella conically rounded, proximally continuing into shaft of volsella. Medial process of volsella set off distinctly from shaft by longitudinal fold (Figs 9, 10).
Description of the male. Colour (Fig. 1): Black, changing into blackish-brown on strongly sclerotized parts of labio-maxillary complex, sterna, coxae, trochanters and femora. The following are light yellowish: large basal spot on mandible; labrum; central area of clypeus except small lateral and ventral margin; broad parallel-sided band at front of frons, shortly interrupted medially on the supra-antennal area, laterally extending obliquely towards upper inner margin of eye where it terminates with straight truncated end; small spot on ocular sinus; narrow streak on tempora along occipital carina at dorso-lateral corner of head; large spot on antero-dorsal angle of pronotum (humeral spot); stripe along dorso-medial (inner) margin of pronotum, posteriorly small becoming distinctly broader antero-medially where it continues into humeral spot in one specimen (Fig. 7); large spot on dorsal mesopleura; tegula except small black marking along anterior part of medial margin; medium-sized median spot on scutellum; propodeal lamella; distal part of femora, tibiae and tarsi (weakly mixed with reddish-brown in parts especially distally); continuous posterior bands on terga I-VI, somewhat smaller on each side of middle (Fig. 6), extending over lateral margin on ventral part of tergum; continuous broad posterior band on sternum II; small spots postero-laterally on sternum III. Reddish-brown are: distal two thirds of mandible; ventral margin of clypeus; claws. Antenna with A1-2 black; A3-6 anteriorly with light yellowish stripe otherwise blackish to dark orange-brown; A7-A12 orange-brown, posteriorly with blackish marking becoming lighter distally, anteriorly and on ventral side markedly orange without black. Wings translucent, weakly yellowish-brown.
Structure: Head in front view as long as broad. Clypeus a little broader than long, distal margin deeply emarginate; cuticula shiny and smooth with close macropunctation becoming less distinct towards distal margin; pale stiff setae arising from macropunctures without distal "knob"; lateral setae with distal ends frequently curved in distal-medial direction. Frons diagonally striated with moderately spaced to close macropunctation, interstices smooth; bearing pale stiff setae, up to 0.22 mm long, 1.1 times as long as diameter of median ocellus, laterally some with curved distal end, in centre mainly with tiny distal "knob"; frontal line in one specimen raised to form a small pro-   tuberance in centre of frons. Median ocellus circular, lateral ocelli more or less obliquely oval; diameter of median ocellus distinctly larger than diameter of lateral ocelli. Vertex falling away obliquely towards occipital carina immediately behind ocelli (Fig. 7); cuticula with close macropunctation becoming more distinct and closely reticulate behind ocelli, where longitudinal interstices are more strongly raised forming lines; cuticula of interstices shiny and smooth; covered with short setae arising from macropunctures. Compound eye sparsely covered with small setae. Gena very narrow; preoccipital carina sharp. Antenna with articles A8-A12 forming ventrally flattened club about 2.0 times as long as broad (viewed dorsally) (Fig. 1); two oval-shaped, perhaps sensory, depressions (tyloids) on concave, ventral side of club, situated within antennal articles A9 and A10.
Anterior margin of pronotum raised to carina especially sharp medially (erroneously termed anterior pronotal carina by Mauss 2013). Short anterior pronotal carina (sensu Carpenter 1988) distinctly present at antero-ventral angle of pronotum running parallel to anterior margin, preceding crenate groove; distance between anterior pronotal carina and anterior margin of pronotum about width of fore metatarsus. Posterior pronotal carina forms narrow translucent sinuate crest on humeral angle of pronotum. Posterior margin of pronotum raised to short carina dorsally in front of tegula. Cuticula of pronotum shiny, with close to reticulate macropunctation, interstices smooth, rounded, with very sparse uneven micropunctation; laterally fairly horizontally striated due to slightly more raised interstices. Cuticula of mesoscutum and mesoscutellum shiny, reticulate with close, deep macropunctation and raised interstices. Postero-medial cuticula of mesoscutum and especially of mesoscutellum with longitudinal interstices becoming narrower and more strongly raised to sharp edges leading to distinctly striated appearance; interstices smooth, laterally on mesoscutellum somewhat finely longitudinally wrinkled. Mesoscutellum laterally with distinct carina along posterior margin, carina medially reduced to a few small tooth-like projections along posterior margin. Carina along posterior margin of metanotum medially with small tooth-like projections. Axilla with short blunt lateral projection only slightly projecting over adjacent, somewhat emarginate, posterior part of tegula. Tegula shiny, closely covered by macropunctures except less densely punctured or completely smooth central convex area. Mesepisternum with pronounced epicnemial carina deflexed backwards to run transversely in front of mid coxa; cuticula shiny, with close macropunctation; horizontally striated by raised interstices; area ventral to scrobal groove coarsely punctured with some interstices strongly raised to knife-like edges forming coarse honeycomb-like sculpture. Process at mesepisternal scrobal groove of moderate size; cuticula on posterior side shiny in centre with irregularly moderately spaced distinct micropunctation, ventrally densely longitudinally wrinkled. Horizontal propodeal triangle laterally delimited by a perpendicular declivity, somewhat laterally produced at postero-lateral edge of propodeal triangle, posteriorly bordered by serrated carina; cuticula shiny, coarsely punctured, interstices almost knife-like. Posterior surface of propodeum striated by strong vertical cuticula-folds; cuticula shiny, without punctuation, weakly coriaceous and covered with short fine pale setae. Cuticula of sides of propodeum and metepisternum shiny, densely horizontally wrinkled. Lateral lamella broad and somewhat convex; lateral margin slightly convex; posterior margin straight, not crenate; medially where lamella joins central part of propodeum with a rounded emargination, ventro-medial edge of which produced to a small blunt protrusion; dorsal cuticula of lamella shiny, with moderately spaced macropunctation, interstices medially smooth, dorso-laterally shagreened. Midcoxa with small but distinct spine at distal end on anterior side close to anterio-medial angle. Claws ventrally with small tooth.
Metasomal terga with dark anterior part continuing into posterior pale part by slight declivity; postero-lateral corners slightly produced; posterior margin of tergum I weakly crenulated, crenulation not produced into spines and not projecting over smooth translucent posterior margin of tergum; posterior margin of terga II-VI crenulated, crenulation produced into short partially posteriorly truncated spines, not or only slightly projecting over smooth translucent lower posterior margin of terga (Fig. 6); cuticula with silken sheen, densely covered with moderately coarse macropunctation, diameter of punctures and distance between them fairly larger on posterior pale part of tergum, with about seven macropunctures along median axis of tergum III; single thin seta arises from bottom of each macropuncture, only slightly protruding over rim of puncture; interstices finely shagreened, moderately covered with very tiny, decumbent, pale setae, all setae orientated towards caudal end. Posterior margin of tergum VII with two lateral deep emarginations, resulting in two lateral lobes that continue laterally into margins of tergum and single medial lobe divided into two lanceolate more or less pointed medial projections of variable length (Fig. 6). Metasomal sternum I shiny, finely shagreened, with tiny setae but without punctures. Sterna II-VI posteriorly with broad stripe of asetose, translucent cuticula adjacent to posterior margin of more strongly sclerotized cuticula; small sparse band of setae along posterior sclerotized margin somewhat projecting over anterior part of translucent stripe of cuticula; sclerotized cuticula shiny, close to anterior margin faintly shagreened, on posterior half of sternum II-VI medially and laterally with sparse punctuation of shallow micropunctures from which short pale setae arise, on sternum II antero-laterally with a few small shallow macropunctures from which short pale setae arise, on sternum III-VI anteriorly with moderate to sparse shallow macropunctation. Sternum VIII with distinct little spine on each side of posterior margin; emargination of posterior margin medially angled, laterally with a transverse section well set of by protruded edge (Fig. 8). Setae only shown on one side of each drawing. Nomenclature follows that of Birket-Smith (1981): ad, aedoeagus; at, apodema thyrsos; cu, cupula; dc, dorso-medial concavity of the posterior margin of the stipes; ha, harpide; mp, medial process of volsella; pp, posterior process of volsella; sl, shovel-like dorso-medial lobe of harpide; sp, stipes; ty, thyrsos; vo, volsella. Male genitalia as in Figs 9-10. Dorso-medial shovel-like lobe of harpide large, densely covered ventrally with long setae. Dorso-posterior margin of stipes with dorso-medial concavity angled. Stipes in dorsal view elongated, with medial and lateral margin running more or less parallel towards posterior end. Sides of stipites converging continuously anteriorly towards cupula. Volsella moderately large and broad, not reaching antero-medial margin of dorso-medial lobe of harpide; dorsal area with strongly sclerotized, large, dark tubercles; distances between tubercles moderate; plane medial process curved more or less hook-like towards posterior end, set off distinctly from shaft by longitudinal fold; posterior process conically rounded, proximally continuing into shaft of volsella. Sides of thyrsoi more or less parallel only slightly converging towards posterior (apical) end of aedoeagus. Each thyrsos continues anteriorly into apodema thyrsos with a distinct outwardly directed curve. On ventral side cupula medially protruded towards anterior end projecting over dorsal margin of cupula in ventral view of genital capsule.
Measurements. Measurements of the exoskeleton are summarized in Table 1. Floral association. Unknown. Distribution. Celonites hermon is known only from three localities in the ranges of the Lebanon and Anti-Lebanon Mountains along the Lebanese section of the Dead Sea Transform (Fig. 11).

Key to separate Celonites hermon from its next relatives C. abbreviatus, C. tauricus and C. andreasmuelleri
For illustrations of the morphological characters of C. abbreviatus, C. tauricus and C. andreasmuelleri see Mauss (2013) and Mauss et al. (2016). Cuticula on frons and at least dorsal half of clypeus densely and strongly shagreened leading to a conspicuous dull region covered with knobbed pollencollecting setae, changing gradually into the adjacent smooth, shiny cuticula on vertex (Fig. 2). Terga of metasoma with yellowish-white to yellow markings without reddish tinge (exception being made for C. abbreviatus from the Peloponnesus with small reddish markings close to the lateral margin of the terga Pollen collecting knobbed setae on frons longer and thinner, up to 1.4 times as long as diameter of median ocellus (Figs 3, 4). Ocelli larger, with diameter of median ocellus distinctly larger than diameter of lateral ocelli (Fig. 4). Lateral ocelli more or less obliquely oval (Fig. 4). Vertex falls away obliquely towards the occipital carina immediately behind the ocelli (Fig. 4) Antennal club (A8-12) with dark tip, that is, at least distal end of A12 is blackish markedly contrasting on ventral side to adjacent light reddish brown area of antennal club. On dorsal side blackish marking extends usually over distal parts of A11 fading gradually towards proximal end (for differences between geographic populations see Mauss et al. 2016 Tergum VII quadrilobed, since the median lobe is medially distinctly emarginate (Fig. 6). Sternum VIII different, with a distinct little spine on each side of the posterior margin (Fig. 8) Posterior margin of stipes of male genitalia with dorso-medial concavity (Figs 9, 10). Stipes in dorsal view elongated, with medial and lateral margin running more or less parallel towards posterior end (Figs 9, 10). Posterior process of volsella conically rounded. Medial process of volsella set off dis-tinctly from shaft by longitudinal fold (Figs 9, 10)

Status and taxonomic position of Celonites hermon
Celonites hermon can be assigned to the subgenus Celonites s.str., the Celonites abbreviatus-group and finally the C. abbreviatus-complex without contradiction as it shares all characters of these taxa established by Richards (1962) and Mauss (2013). Among the members of the C. abbreviatus-complex Celonites hermon resembles C. andreasmuelleri most of all in the comparatively thin pollen collecting knobbed setae that are of equal diameter along most of their length, the enlarged ocelli, with a distinctly large circular median ocellus and smaller oval shaped lateral ocelli and the characteristic form of the head, that falls away obliquely towards the occipital carina immediately behind the ocelli. Moreover, in the male genitalia the posterior margin of the stipes has a characteristic dorso-medial concavity and the medial and lateral margin of the stipes are running more or less parallel towards the posterior end. These distinctive similarities are probably synapomorphic characters of both taxa, as they are not present in other members of the C. abbreviatus-complex and even the whole C. abbreviatus-group.
On the other hand, Celonites hermon is lacking some of the potential autapomorphies of C. andreasmuelleri recognized by Mauss (2013), i.e. the characteristic semi-circular cuticula structure on the frons of the females, as well as the differing form of sternum VIII and tergum VII and the considerably larger shovel-like dorso-medial lobe of the harpide in the males of C. andreasmuelleri. Moreover, both taxa differ in colour, the colour pattern on the ventral side of the tip of the antennal club and the form of the volsella. The distinct differences in the male genitalia, the form of sternum VIII and tergum VII as well as the colour pattern on the ventral side of the antennal club that has been assumed by Mauss et al. (2016) to be associated with mate recognition, lead to the conclusion that both taxa are probably reproductively isolated and represent two separate biospecies (sensu Mayr 1967).
The hypotheses proposed by Mauss (2013) that Celonites hermon is identical with C. abbreviatus, is rejected. Both species differ clearly in form and length of the pollen collecting knobbed setae, the size of the ocelli and the shape of the vertex. Moreover, there are distinct differences in the male genitalia. The misinterpretation of the morphological characters of the type by Mauss (2013) resulted to some extend from the mediocre condition of the female specimen, of which the setae on frons and vertex are stuck together to a high degree by remnants of a viscous liquid, giving them a shorter and stouter appearance and partly covering the exoskeleton.

Biogeography
Celonites hermon has been recorded only from a small area in the mountain ranges along by both sides of the Lebanese section of the Dead Sea Transform (Fig. 11). Since C. hermon differs significantly from C. abbreviatus in various morphological characters, it cannot represent a relict population of C. abbreviatus as implied by Mauss (2013). Moreover, the morphological similarities between C. hermon and C. andreasmuelleri indicate a close relationship between them and a more distant relationship to C. abbreviatus and C. tauricus. Therefore, it can be hypothesized that C. hermon and C. andreasmuelleri originated from a common stem species that inhabited the area along the Dead Sea Transform. Geographic isolation and succeeding speciation may have led to the recent allopatric distribution with C. hermon being restricted to the northern Lebanese section of the Dead Sea Transform and C. andreasmuelleri to the southerly adjacent area of the Jordan Rift Valley (Fig. 11).