Holaptilon brevipugilis Kolnegari, 2018

Mirzaee, Zohreh, Battiston, Roberto, Ballarin, Francesco, Sadeghi, Saber, Simoes, Marianna, Wiemers, Martin & Schmitt, Thomas, 2024, The Six Dwarfs of the Middle East: revision of the enigmatic praying mantis genus Holaptilon (Mantodea: Gonypetidae: Gonypetinae) with the description of four new species under integrative taxonomy, Arthropod Systematics & amp; Phylogeny 82, pp. 89-117 : 89

4.5. Holaptilon brevipugilis Kolnegari, 2018

Figure 9i, j

H. yagmur Yılmaz and Sevgili, 2023: 18. new. syn.

Material examined.

Paratypes: 1♂, 1♀, ethanol, with genitalia in a separate micro-vial, Arak , Markazi province, Iran, 34.128N, 50.072E, 1803 m. 7/2018, leg. Kolnegari (SDEI) . - Other material: 1♀ ethanol, Arak , Markazi province, Iran, 34.128N, 50.072E, 1803 m . 7/2018, leg. Kolnegari (SDEI); 1♂, ethanol, Mashhad , Khorasan Razavi, Iran, 36.316N, 59.410E, 995 m . 8/2021, leg. Ghafarnia (ZMPC), 1 nymph, Siverek , Karabahçe, Turkey, 37.776N, 39.735E , 08/2019, leg. K. Yılmaz and M. Yalçın (HSPC).

New diagnosis.

Short afa, which is finger-like, less markedly narrowed towards apex, curved; pba anteriad of afa less sclerotised; posterior edge of vla oblique (Fig. 2g, h ).

Redescription.

Males are smaller and more delicate in appearance than females, both male and female apterous, head and body dorsally sandy brown, with small dark brown and black spots, mostly in the middle of body parts. Head: Wider than high (ratio: 1.2-1.3), wider (ratio: 1.1) than pronotum (Table S3, Fig. 10i-l ). Pronotum: Rectangular shape, almost flat, compact. Slightly higher than wide (ratio: 1.1) Meso- and metanotum: roof-shaped and slightly keeled (Fig. 11i-l ). Coxae robust, shiny blue-black in some of the individuals but in some others not shiny black or only 1/3 black, anterior and posterior margin with some tubercles and setae. Femora broad, dorsal edge lamellar, two times longer than wide, armed with 10-12 anteroventral spines with the second one longer than the others; 4 discoidal spines with the first one shorter, the third one longer than the others, the second one is a bit smaller than the third one, the forth is short but longer than the first one; 4 posteroventral spines, with the first one slightly longer than the other three spines, the first two spines are close to each other but the third and fourth have a bit more distance between them; anterior genicular lobe and posterior genicular lobe with a spine; foretibia armed with 9 anteroventral spines, elongating distally, and 10-13 posteroventral spines, also elongating distally (Fig. 12c ). Abdomen: Slender but half as wide in male compared to female, the tergites keeled in midline; supra anal plate transverse, triangular; cerci with eight readily recognizable cercomeres, covered by setae; last cercomere longer and narrower than the others; subgenital plate much longer than wide. Male genitalia: Ventral phallomere oval, moderately wide; afa short, which is finger-like, less markedly narrowed towards apex, curved; pba anteriad of afa less sclerotised. posterior edge of vla oblique; sclerite L4B curved, of complex shape, widened distally. Apical process paa long, directed left side, with curved apex (Fig. 2g, h ). Ootheca: Dark yellow to creamy in colour, oval-shaped in dorsal view, dorsoventrally flattened. Rows of egg chambers that are parallel to the top and bottom of the egg, casing is visible from outside (Kolnegari and Vafaei 2018).

Distribution.

Centre and north-east of Iran, Markazi province (Arak) (Kolnegari and Vafaei 2018), eastern Turkey ( Yılmaz and Sevgili 2023), (Fig. 6 : purple points).

We record this species for the first time from Khorasan Razavi, Mashhad, Iran.

Conservation.

This species is widely distributed with an Extent of Occurrence (EOO) of about 290.000 km2 in an apparently very fragmented number of locations (3) but this number is probably underestimating the real number of locations. With no data on population trends over time, low anthropogenic presence and impacts in its known localities, this species can be addressed as Least Concern. Further research is however encouraged to collect more data and information on its real distribution and threats.

Remarks.

The two paratypes of H. brevipugilis we were able to study and compare with our own material were collected at the same locality and on the same date as the holotype, so we are confident that they represent the same taxon. The original description of this species (Kolnegari 2018) was based on a very small number of specimens and on qualitatively variable morphological characters which were found to be present also in other species. A new species diagnosis is therefore proposed above, based on male genitalia here described for the first time.

The taxonomic status of Holaptilon yagmur underwent re-evaluation after determining its phylogenetic position within the genus. Our molecular genetic investigation revealed that the conventional reliance on external morphological traits for differentiating H. yagmur from other conspecifics within the genus lacked effectiveness in achieving precise species identification. Through comprehensive analysis of two mitochondrial (COI, 16S rDNA) and two nuclear DNA markers (H3, IDH), our molecular data showed that H. yagmur and H. brevipugilis are genetically indistinguishable or very closely related (Fig. 4 ). Consequently, H. yagmur should be considered synonymous with H. brevipugilis .

Figures

Figure 2. Dorsal view and ventral view of male genitalia of different Holaptilon species: a, b H. abdullahii sp. nov.; c, d H. iranicum sp. nov.; e, f ventral view H. tadovaniensis sp. nov.; g, h H. brevipugilis; i, j H. khozestani sp. nov.; k, l H. pusillulum; scale bar: 500 µm. afa = anterior process (left phallic complex: left phallomere). paa = apical process (left phallic complex: left phallomere). vla = the right-posterior ventral lobe (left phallic complex: " ventral phallomere "; with L 4 A sclerotization in ventral wall; with opening of ejaculatory duct in dorsal wall). pba = (left phallic complex: left phallomere), of afa plus the edge (between pouches pne and lve) from which they arise (with L 1 and L 2 sclerotizations). L 4 B = sclerite extending over the dorsal wall (left phallomere). L 4 A = sclerite extending over the ventral wall (left and " ventral " phallomere).

Figure 4. Phylogenetic trees of Holaptilon: a Phylogeny and Molecular Species Delimitation of the genus Holaptilon using a combined phylogenetic tree made through Bayesian (BI) and Maximum Likelihood (ML) approaches with the utilization of MrBayes and IQ tree, respectively (both trees did not differ in their topology) based on concatenated COI, 16 S rRNA, H 3, and IDH sequences, showing the phylogenetic placements of the new samples from Iran and results of three molecular species delimitation methods, ASAP: Assemble Species by Automatic Partitioning; bPTP: Bayesian implementation of the PTP model for species delimitation; mPTP: multiple-rate Poisson Tree Process. Support values are indicated beside the nodes [BI posterior probability / ML bootstrap (PP / BP)], holotypes are marked with asterisks (*); con: concatenated, nu: nuDNA. The BI tree was used as the basis for this figure. b Phylogenetic tree made using Bayesian (BI) method of the genus Holaptilon based on concatenated COI and 16 S rRNA sequences; c) Phylogenetic tree made using Bayesian (BI) method of the genus Holaptilon based on concatenated H 3 and IDH sequences.

Figure 6. Current and historic records of Holaptilon species, visualisation with QGIS v. 3.22, Base map: ESRI Terrain map.

Figure 9. Genus Holaptilon life habitus: a H. abdullahii sp. nov., paratype male from Soroo, Busheher province (29.569 N, 51.947 E). b H. abdullahii sp. nov., paratype female from Kangan, Busheher province (27.843 N, 52.064 E). c H. brevipugilis male and female from Arak (34.128 N, 50.07 E) (photo credit: Mahmood Kolnegari). d H. iranicum sp. nov., holotype male from Arjan, Fars province (29.569 N, 51.947 E). e H. iranicum sp. nov., paratype female from Arjan, Fars province (29.569 N, 51.947 E). f H. khozestani sp. nov., holotype male from Malagha, Khozestan province (31.607 N, 49.998 E). g H. khozestani sp. nov., paratype female from Dehdez, Khozestan province (31.733 N, 50.222 E). h H. tadovaniensis sp. nov., paratype female from Tadovan, Fars (28.853 N, 53.326 E). i H. tadovaniensis sp. nov., holotype male from Tadovan, Fars (28.853 N, 53.326 E). j H. pusillulum male from Jerusalem, Israel (28.853 N, 53.326 E) (photo credit: More Yosef Avi). k H. pusillulum female from Jerusalem, Israel (31.737 N, 35.077 E) (photo credit: Chaym Turak).

Figure 10. Heads of different Holaptilon species: a male (Kangan), b female (Kangan) H. abdullahii sp. nov.; c female (Arjan), d male (Arjan), e female (Sahlak), f female (Yasuj) H. iranicum sp. nov.; g male (Tadovan), h female (Tadovan) H. tadovaniensis sp. nov.; i female (Arak), j male (Mashhad), k female (Arak), l male (Arak) H. brevipugilis; m male (Dehdez), n male (Malagha), o male (Dehdez), p female (Dehdez) H. khozestani sp. nov.; scale bar: 1 mm.

Figure 11. Posterior view of the thoraxes of different Holaptilon species: a male (Kangan) b female (Kangan) H. abdullahii sp. nov.; c female (Arjan), d male (Arjan), e female (Sahlak), f female (Yasuj) H. iranicum sp. nov.; g male (Tadovan), h female (Tadovan) H. tadovaniensis sp. nov.; i female (Arak), j male (Mashhad), k female (Arak), l male (Arak) H. brevipugilis; m male (Dehdez), n male (Malagha), o male (Dehdez), p female (Dehdez) H. khozestani sp. nov.; scale bar: 2 mm; white arrows indicate the characters that were used in the previous studies as diagnostic characters.

Figure 12. Different types of spines of the foreleg of different Holaptilon species: a female H. iranicum sp. nov.; b female H. tadovaniensis sp. nov.; c male H. brevipugilis; d male H. khozestani sp. nov.; e male H. abdullahii sp. nov.; f female H. pusillulum; scale bar: 2 mm.