Histopona leonardoi, Bolzern, Angelo, Pantini, Paolo & Isaia, Marco, 2013

Bolzern, Angelo, Pantini, Paolo & Isaia, Marco, 2013, Revision of the Histopona italica group (Araneae: Agelenidae), with the description of two new species, Zootaxa 3640 (1), pp. 23-41 : 33-38

Histopona leonardoi sp. n.

Figures 7–8 , 12 , 17–20 , 25–26 , 29

H. italica Brignoli, 1977: 35 , f. 14–15, (m misidentified).

Type material. Holotype male: ITALY: Piemonte, Cuneo: Acceglio, springs of Maira River, 1680 m, UTMED50 32T 335892 4926442 (lon. 6.9366° lat. 44.4726°), sparse larch wood, 3 4/VI/2009, Rosso M.

Paratypes: ITALY: Val d'Aosta: Aosta: Ayas, Champoluc, 1700 m, UTMED50 32T 401771 5076342 (lon. 7.7352° lat. 45.8336°), sparse larch wood, 13 31/VIII/2007, 13 15/VII/2009, Franco I. (CI); Gressoney-St. Jean, 2100 m, UTMED50 32T 409050 5067251 (lon. 7.8306° lat. 45.7528°), alpine praires 13 7/IX/2007, Negro M. (CI, Negro et al. 2009 sub H. italica ); Gressoney-La-Trinité, 1700 m, UTMED50 32T 407631 5079494 (lon. 7.8100° lat. 45.8628°), sparse larch wood, 13, 4Ƥ 30/VI/2006, Negro M. (NMB: 20853, Negro et al. 2009 sub H. italica ); Gressoney-La-Trinité, Gabiet, 2458 m, UTMED50 32T 410779 5079371 (lon. 7.8506° lat. 45.8621°), alpine praires, 2Ƥ 20/VIII/2008, Negro M. (CI, Negro et al. 2010 sub H. italica ). Piemonte: Moncerchio di Vallanzengo, Val Sessera, UTMED50 32T 428560 5058143 (lon. 8.0827° lat. 45.6731°), beech wood, 3 m, 2Ƥ 2/V/2009, 63 5/ IX/2009, 583, 1Ƥ 2/IX/2009 Franco I., Negro M.; 2Ƥ 2/V/2009, Franco I.; Cuneo: Acceglio, springs of Maira River, 1680 m, UTMED50 32T 335892 4926442 (lon. 6.9366° lat. 44.4726°), sparse larch wood, 1Ƥ 4/VI/2009, Rosso M. (CI); Entracque, Natural Park of Alpi Marittime, 1100m, UTMED50 32T 376018 4893859 (lon. 7.4487° lat. 44.1874°), beech wood close to Busset stream, 83, 5 Ƥ 29/VI–9/VIII/2007, Wolf-Schwenninger, 2Ƥ 21/IX/ 2008, Isaia M., Paschetta M. (CI); Terme di Valdieri, Natural Park of Alpi Marittime, Vallone del Valasco, UTMED50 32T 358295 4895033 (lon. 7.2267° lat. 44.1947°), alpine pasture with sparse larch wood, 73, 1Ƥ 11/ VII–27/VIII/2009, Isaia M., Paschetta M.; Natural Park of Alpi Marittime, Pian della Casa, 1473 m, UTMED50 32T 361775 4896277 (lon. 7.2699° lat. 44.2066°), alpine pasture, 13 11/VII/2008, Isaia M., Paschetta M. (CI, Paschetta et al., 2012 sub H. italica ); Natural Park of Alpi Marittime, Piano del Valasco, UTMED50 32T 358300 4895039 (lon. 7.2267° lat. 44.1948°), alpine pasture with sparse larch wood, 83, 2Ƥ 27/VIII–23/IX/2009, Isaia M., Paschetta M. (MSNVR); Terme di Valdieri, Natural Park of Alpi Marittime, 1368 m, UTMED50 32T 362064 4896332 (lon. 7.2735° lat. 44.2071°), beech wood, 1Ƥ 29/VI/ 2009, Isaia M., Paschetta M. (CI); Vernante, Natural Park of Alpi Marittime, Palanfrè, 1370 m, UTMED50 32T 379547 4894524 (lon. 7.4926° lat. 44.1939°), beech wood, 1Ƥ 10/IX/2008, 2Ƥ 22/VI/2009, Isaia M., Paschetta M., 2Ƥ 2/VII/2010, Isaia M. (CI); Torino: Ribordone, Santuario Prascundù, 1400 m, 1Ƥ 28/IX/2004, Giachino P.M.; cave “Tuna del Diau, 1621 Pi/TO”, 1080 m, UTMED50 32T 350672 4979013 (lon. 7.1070° lat. 44.9488°), 13 5/X/2002, Lana E. (CI, Isaia et al., 2011 sub H. italica ); Vistrorio, 1Ƥ V– VIII/1993, Giachino P.M.; Verbania-Cusio-Ossola: Varzo, cave “Grotta di San Carlo”, 1Ƥ 4/VI/1978, Casale A. (MSNVR, Brignoli, 1979 sub H. italica ). Liguria: Genova: Mezzanego, Ghiaiette, UTMED50 32T 537328 4918352 (lon. 9.4688° lat. 44.4174°), beech wood 850 m, 2Ƥ 31/X/2009 – 25/V/2010, 13 25/V–18/VIII/2010 Lodovici O., Pantini P., Valle M.; Mezzanego, Forest of Monte Zatta c/o ex Colonia Devoto, 1050 m, UTMED50 32T 535942 4917017 (lon. 9.4513° lat. 44.4055°), beech wood, 2Ƥ 31/X/2009 – 25/V/2010, Lodovici O., Pantini P., Valle M., 1Ƥ 25/V/2010, 53, 7Ƥ 25/V–18/VIII/2010 Lodovici O., Pantini P.; Propata, north slope of Monte Cremado, 1640 m, 1Ƥ 5/VI–12/VII/1988, Cartasegna F., Pesce D. (CG); Torriglia, Passo del Colletto, 1280 m, 1Ƥ 21/V–1/VII/1999, Pesce D. (CG); Torriglia, SE slope of Monte Duso, 1380 m, 1Ƥ 21/V–1/ VII/1999, Cartasegna F. (CG); La Spezia: Varese Ligure, Passo Cento Croci, UTMED50 32T 549768 4918763 (lon. 9.6251° lat. 44.4204°), 1000 m, 43, 1Ƥ IV– VIII/1991, Cerbino R., Valle M., 23 VI–IX/1992, Pantini P., Valle M.; Savona: Sassello, Rio del Nido, 1000 m, beech wood, 43, 18/VII–10/X/2001; Sassello, Monte Beigua, 1000 m, 1Ƥ 17/VII/2001. Lombardia: Pavia: Santa Margherita di Staffora, Hotel Colletta, 1380 m, beech wood, 113 31/VII–19/IX/2001, 43, 7Ƥ 19/IX/2001 – 20/III/2002, 1Ƥ 26/IV–27/VI/2002, Pantini P. Emilia Romagna: Parma: Bedonia, Passo di Montevacà, 800 m, UTMED50 32T 548856 4931317 (lon. 9.6149° lat. 44.5335°), 13 IX/1991 – V/1992, Buttarelli G., Cerbino R., Pantini P., Valle M.; Piacenza: Bobbio, Passo Penice, 1100 m, UTMED50 32T 525987 4960430 (lon. 9.3285° lat. 44.7967°), wood, 6Ƥ 20/V–20/VI/2001 (3Ƥ NMB: 20536), 1Ƥ 20/VI–31/VII/2001, 13 31/VII–19/IX/2001, 23, 4Ƥ 19/IX/2001 – 20/III/2002, 1Ƥ 20/III–26/IV/2002 Pantini P.; Bobbio, road to Monte Penice, 1400 m, UTMED50 32T 525246 4959349 (lon. 9.3191° lat. 44.7870°), wood, 133, 1Ƥ 31/VII–19/IX/2001, road margin 1400 m, 53, 1Ƥ 19/IX/2001 – 20/III/2002, 1Ƥ 26/IV–27/VI/2002 Pantini P.

Other material examined. SWITZERLAND: Tessin: Centovalli, Lionza, 930 m, UTMED50 32T 470518 5112649 (lon. 8.6181° lat. 46.1667°), 13, 2Ƥ, 6/VI/1989, 5/VII/1989, 11–25/VIII/1989, Hänggi A. (NMB: 0 2488 d, 20671-20672; Hänggi 1992, sub H. italica ); Val Careccio, 23, 1Ƥ, 29/IV–19/IX/1988, Pronini, P. (NMB: 20669- 20670; Pronini 1989 sub H. italica ). ITALY: Piemonte: Biella: Oropa, 13 24/VIII/ 1972, Vigna Taglianti A. (MSNVR, paratype of H. italica , misidentification); Crissolo, Monviso, 1300 m, 1Ƥ VII/1967, Osella G. (MSNVR, paratype of H. italica , misidentification).

Etymology. The species is dedicated to Leonardo Pantini, firstborn of PP. The species epithet is a name in apposition.

Diagnosis. Males ( Figures 7–8 , 17–20 ) can be separated by the absence of a patellar apophysis (present in torpida group, except H. vignai Brignoli 1980 ), the distally spoon-like elongated radix (absent in myops - and strinatii group, plate-like and distally bifid in H. italica , tube-like in H. fioni sp. n.) and the distally broadly rounded conductor (strongly elongated in H. fioni sp. n.). Females ( Figures 25–26 , 29 ) can be separated from other Histopona species by the glossy median indented posterior epigynal sclerite (much longer and with anterior margin only moderately indented in torpida group) with moderately diverging margin (parallel in H. italica , strongly diverging in H. fioni sp. n.), the unpaired “bursa copulatrix” (completely paired copulatory ducts in myops - and strinatii group) with anterior margin concave (straight or convex in H. italica , v-shaped in H. fioni sp. n.) and the narrow lateral lobes of the copulatory ducts (broad in H. italica ). See also Table 1 .

Description. Measurements and ratios of male (n=2, holotype and paratype from Entracque): carapace 2.25– 2.86 long, 1.54–2.05 wide. Head region 0.80–1.10 wide; PER 0.45–0.62 wide. Chelicerae 1.02–1.34 long, 0.46– 0.56 wide. Labium as long as wide. Gnathocoxa ratio width to length: 0.508–0.543. Sternum 1.23–1.51 long, 1.05– 1.25 wide. Opisthosoma 1.98–2.46 long, 1.00–1.35 wide. Ratio bulb length (laterally from cymbium base to conductor tip) to cymbium length: 0.670–0.749. Leg measurements are given in Table 2.

Measurements of females (n=2, Paratype females from Acceglio and Entracque): carapace 2.04–2.28 long, 1.38–1.63 wide. Head region 0.81–1.01 wide; PER 0.48–0.54 wide. Chelicerae 0.87–1.04 long, 0.45–0.49 wide. Labium as long as wide. Gnathocoxa ratio width to length: 0.56. Sternum 1.20–1.25 long, 1.00–1.08 wide. Opisthosoma 2.01–2.69 long, 1.35–1.81 wide. Epigynal plate 0.70–0.72 long, 0.76–0.78 wide; atrium 0.16–0.18 long, 0.67–0.70 wide. Leg measurements are given in Table 2.

Eyes: in dorsal view both eye rows straight or slightly recurved; in frontal view AER straight or slightly procurved, PER procurved. Diameters: PME: 0.103–0.128; PLE: 0.096–0.129; AME: 0.059–0.082; ALE; 0.109– 0.130. Distances: PME–PME about half diameter of PME or slightly less; PME–AME about half diameter of PME or slightly less; PME–PLE about half diameter of PME; PME–ALE about half diameter of PME or slightly less; AME–AME about half diameter of AME or slightly less; AME–ALE less than half diameter of AME. Clypeus height (measured under AME) about 3 diameters of AME or slightly more; clypeus height (measured under ALE) about twice diameter of ALE or slightly less.

Coloration: Carapace with narrow, continuous dark margin; two longitudinal symmetric darkened rows of triangular dots present on carapace; narrow darkened band median at head region present. Sternum without pattern. Opisthosoma dark grey green; cardiac mark moderately pronounced; posteriorly with indistinct pattern of chevrons. Legs without color pattern.

Additional somatic characters: distal margin of labium weakly concave. Plumose hairs present on carapace, legs and opisthosoma. Three promarginal teeth, the second one from proximal biggest; 5–7 retromarginal teeth, all equal in size. All trochanters notched. All tarsi with 6–7 dorsal trichobothria. No trichobothria on palp tarsi or cymbium. Pale colulus, sometimes moderately darkened, divided into two plates. PLS longer than all others with distal segment as long as basal segment, both moderately darkened. PMS as long as ALS. ALS moderately darkened. The formulae of leg spination are listed in Table 3 .

Male palp ( Figures 7–8 , 12 , 17–20 ): RTA with a big dorsal branch, distally pointed, strongly sclerotized and moderately stepped; lateral branch forming moderately sclerotized finger-shaped appendix; ventral branch forming bulge-like moderately ventrodistally protruding stepped appendix, lateral with 2–3 small stepped bands. Tegulum broadly ring-shaped, distally dividing into a filiform embolus and an elongated, distally spoon-like apophysis (radix), terminally often with a transparent portion. Embolus originating (free apex) at 11 o'clock position; distal tip between 3 and 4 o’clock position. Conductor lamella-like, distally broadly rounded and moderately elongated, laterally folded along the whole length; shorter than alveolus, distally not reaching over alveolus margin; terminal end forming moderately sclerotized peak. Connection of conductor and tegulum membranous, band-like. Median apophysis and tegular apophysis absent.

Epigynum and vulva ( Figures 25–26 , 29 ): rectangular epigynal plate sclerotized, posterior with distinct atrium; atrium anteriorly limited by strongly sclerotized, m-shaped margin of the epigynal plate with a posteriorly tapered median region; atrium posteriorly limited by a glossy sclerite (“epigynal valve”), median deeply indented with diverging margins; between anterior margin and posterior sclerite atrium covered by membranous white cuticula. Copulatory openings located at anteriolateral border of atrium. Copulatory duct first unpaired (“bursa copulatrix”), then dividing into narrow paired lateral lobes directing into strongly sclerotized convoluted receptacula; fertilization ducts very short.

Distribution. Italy and Switzerland (Tessin). All along the Western Alps and the Northern Apennine.

Ecology. Records of H. leonardoi sp. n. mostly refer to forest habitats (beech woods at an elevation of 1000– 1500 m). The species also occurs at higher elevation in alpine pastures (maximum elevation reached at 2458 m Gabiet, Aosta Valley). In a few cases H. leonardoi sp. n. occurred in caves. Adults are preferably found from spring to autumn.

Anterior limitation of atrium M-shaped margin of the Almost straight margin M-shaped margin of the

epigynal plate with a posteriad of the epigynal plate epigynal plate with a

tapered median region posteriad tapered median

vulva region and Median posterior margins sclerite of glossy Almost parallel Strongly divergent Divergent Epigynum Shape Figures copulatory of 27 anterior – duct 29) (arrows part of in Straight or moderately convex Concave, v-shaped Concave

Lateral lobes of copulatory Very broad, distinct Narrow, band-like Narrow, band-like ducts

Histopona . italica Brignoli 1977

Paratype male from Apecchio

fe pa ti mt ta total Palp 1.15 0.49 0.40 - 1.50 3.54 I 2.28 0.93 2.01 2.14 1.61 8.97 II 2.06 0.89 1.55 1.83 1.27 7.60 III 2.04 0.84 1.58 2.04 1.14 7.64 IV 2.68 1.01 2.26 2.94 1.43 10.32 Paratype female from Apecchio

Palp 0.95 0.46 0.65 - 1.11 3.17 I 2.03 0.88 1.66 1.72 1.41 7.70 II 1.86 0.85 1.36 1.51 1.08 6.66 III 1.85 0.75 1.34 1.81 1.08 6.83 IV 2.30 0.88 2.01 2.60 1.40 9.19 H. fioni sp. n.

Holotype male and paratype male from Pagnona (n=2)

Palp 1.23–1.40 0.49–0.58 0.43–0.48 - 1.54–1.79 3.69–4.25 I 2.61–2.79 1.00–1.03 2.36–2.52 2.42–2.61 1.82–1.97 10.21–10.92 II 2.45–2.72 0.97–1.06 1.97–2.12 2.27–2.42 1.64–1.82 9.3–10.14 III 2.42 0.94 1.85 2.45 1.52 9.18 IV 3.06–3.33 0.97–1.00 2.67–2.85 3.42–3.64 1.85–1.97 11.97–12.79 Paratype females from Pagnona and Rovereto (n=2)

Palp 1.18–1.23 0.49–0.60 0.51–0.85 - 1.05–1.26 3.23–3.94 I 2.64–2.75 1.09–1.15 2.30–2.50 2.24–2.50 1.67–1.85 9.94–10.75 II 2.45–2.60 1.00–1.06 1.94–2.10 2.15–2.25 1.58–1.60 9.12–9.61 III 2.36–2.42 0.94–0.96 1.82–1.96 2.31–2.36 1.30–1.31 8.73–9.01 IV 2.97–3.08 1.04–1.06 2.64–2.77 3.21–3.35 1.67–1.73 11.53–11.99 H. leonardoi sp. n.

Holotype male and paratype male from Entracque (n=2)

Palp 0.89–1.19 0.38–0.47 0.33–0.38 - 1.01–1.63 2.61–3.67 I 2.07–2.33 0.76–0.91 1.81–2.12 1.92–2.15 1.34–1.63 7.90–9.14 II 1.95–2.25 0.78–0.86 1.51–1.77 1.78–1.99 1.26–1.52 7.28–8.39 III 1.86–2.06 0.69–0.93 1.40–1.69 1.91–2.25 1.11–1.24 6.97–8.17 IV 2.42–2.73 0.78–0.92 2.09–2.42 2.71–3.15 1.44–1.58 9.44–10.8 Paratype females from Acceglio and Entracque (n=2)

Palp 0.78–0.84 0.32–0.37 0.50–0.52 - 0.88–0.91 2.48–2.64 I 1.70–1.88 0.70–0.83 1.38–1.61 1.40–1.64 0.92–1.28 6.10–7.24 II 1.52–1.78 0.68–0.73 1.10–1.23 1.32–1.48 0.94–1.06 5.56–6.28 III 1.50–1.72 0.66–0.73 1.08–1.29 1.38–1.68 0.76–0.90 5.38–6.32 IV 1.94–2.25 0.74–0.79 1.62–1.92 2.04–2.45 1.02–1.19 7.36–8.60 According to the identification key provided by Deeleman-Reinhold (1983), Histopona italica forms a singlespecies group within the genus. The two new species described in this work increase the membership of the italica group, which is defined for females by the presence of a glossy, median deeply indented posterior epigynal sclerite and by the unpaired copulatory ducts, and for males by the absence of a patellar apophysis and by the shape of the embolus, originating basal to the protruding radix.

During the examination of the material presented here, large differences in the size of the male palp could be observed, even between specimens from the same locality (e.g. 23 from Liguria, La Spezia: Varese Ligure, Passo Cento Croci). Within the examined specimens of H. leonardoi sp. n., two males appeared to be distinctly larger, the palps showing minor morphological differences ( Figures 17–18 ). Likewise, Bolzern et al. (accepted) and Kraus (1955) observed high intraspecific variation in different species of agelenids, like Malthonica picta (Simon 1870) and Tegenaria femoralis Simon 1873 . The absence of valid morphological characters (the only significant difference being the size) and the lack of concrete evidence (no different female forms found and syntopy of both male forms - same locality and same time of the year) provide convincing support to consider the larger males as larger forms of the same species.

Records of species belonging to the Histopona italica group are known from large parts of Italy (from Calabria to Trentino, along the entire Appenine range, through the Western Alps up to the Lombardian Prealps) ( Figure 30). In some cases, specimens of H. italica and H. leonardoi sp. n. were collected at the same locality. Accordingly, the known distribution of H. italica overlaps that of H. leonardoi sp. n. in the district of Maritime Alps and Northern Apennines, the first extending southwards along the Apennines and the latter northwards, along the Alps. It is likely that H. leonardoi sp. n. and H. italica may also occur in French Maritime Alps, but no direct evidence support the presence of this species in France.

Records of H. fioni sp. n. are only known from the Lombardian Prealps (Lombardia and southern Trentino in Italy and Tessin in Switzerland). Apparently, no overlap occurs between H. fioni sp. n. and H. leonardoi sp. n., being separated by Lake Maggiore, at the border between Piemonte and Lombardia (Tessin Valley). The same separation is known to occur for Coelotes pickardi tirolensis (Kulczyn'ski 1906) and C. pickardi pickardi O. P.- Cambridge 1873 (Isaia & Pantini 2009) and for Troglohyphantes lucifuga Simon 1884 and T. sciakyi Pesarini 1989 (Isaia & Pantini 2010) .

Figures

FIGURES 1 – 8. Left male palp in ventral, dorsal, dorsolateral and retrolateral view. 1, 2. Histopona italica Brignoli 1977 (paratype, Marche); 3, 4, 5, 6. H. fioni sp. n.; 7, 8. H. leonardoi sp. n. C: conductor; E: embolus; R: radix; RTA: retrolateral tibial apophyses; RTAd: dorsal branch of RTA; RTAl: lateral branch of RTA; RTAv: ventral branch of RTA.

FIGURES 11 – 12. Chelicerae and tibia with RTA of left male palp in dorsoretrolateral view. 11. Histopona fioni sp. n. 12. H. leonardoi sp. n. RTA: retrolateral tibial apophyses; RTAd: dorsal branch of RTA; RTAl: lateral branch of RTA; RTAv: ventral branch of RTA. Scale = 1.0 mm (11) and 0.5 mm (12).

FIGURES 13 – 20. Left male palp in ventral, and retrolateral view. 13, 14. Histopona italica; 15, 16. H. fioni sp. n.; 17 – 20. H. leonardoi sp. n., large sclerotized (17 – 18) and small “ normal ” palps (19 – 20). Scale = 1.0 mm. R: radix. The arrow indicates the distinctly stepped connection between conductor and tegulum in H. italica.

FIGURES 21 – 26. Epigyne and vulva in ventral view. Vulva cleared with clove oil. 21, 22. Histopona italica (holotype); 23, 24. H. fioni sp. n.; 25, 26. H. leonardoi sp. n. Scale = 1.0 mm. EV: epigynal valve.

FIGURES 27 – 29. Schematic drawing of vulva in ventral view. 27. Histopona italica; 28. H. fioni sp. n.; 29. H. leonardoi sp. n. CO: copulatory opening; EV: epigynal valve; FD: fertilization duct; latCD: lateral lobe of the copulatory duct; RC: receptaculum; Arrows indicate the posterior margin of the copulatory duct. Scale = 1.0 mm.

Tables

TABLE 1. Diagnostic characters for Histopona italica Brignoli 1977, H. fioni sp. n. and H. leonardoi sp. n.

Character	H. italica Brignoli 1977	H. fioni sp. n.	H. leonardoi sp. n.
Ventral branch of retrolateral tibial apophysis (RTAv)	Strong protruding, distally rounded	Moderately protruding, distally stepped	Moderately protruding, distally with 2–3 small stepped bands
Radix palp Male Conductor	One pointed and one rounded, plate-like end Distally broad rounded and moderately elongated	Tube-like Distally strongly elongated	Plate or spoon-like Distally broad rounded and moderately elongated
Connection conductor-tegulum	Distinctly stepped (arrow in Figure 14)	Continuous	Continuous

TABLE 3. Spination of legs of Histopona italica Brignoli 1977, H. fioni sp. n., H. leonardoi sp. n. The formula gives the number of spines as follows: dorsal - prolateral - retrolateral - ventral; p indicates that the spine is paired (1 p = 2 spines); s indicates the presence of a short and strong spine. A superscript “ - “ or “ + ” indicates that a lower or a higher number of spines have been occasionally observed at this position.

Leg Species	fe	pa	ti	mt	ta
Palp H. italica	3–0–0–0	2–0–0	1+–2–0–0	-	-
H. fioni	2+–0–0–0	2–0–0	1+–2-–0–0 2–1+1p–0–0 2–2p–0–0	-	-
H. leonardoi	2–0–0–0	2–0–0	1+–2-–0–0	-	-
Leg I H. italica	3+–1+–1–0	2–0–0	2–1–0–3p 2–2–0–1+2p	0–0–0–1+2p+1 0–1–0–3p+1	0
H. fioni	1++–1+–0+–0	2–0–0	2--–1+–0–3p 2–2–0–2p+1	0–0+–0–3p+1	0
H. leonardoi	1++–1–0+–0	2–0–0	2–1–0–1p 2–2–0–1+1p+1 2–2–0–1+2p 2–2–0–1p+ +1	0–0–0–3p+1	0
Leg II H. italica	2++–1–1–0	2–0–0	1+–2-–0–1+1p+	0–2–0++–1+2p+1 0–4–1–1+2p+1	0
H. fioni	3–1++–1+–0	2–0–0	2–2–0+–3p	0–2–0+–3p+1	0
H. leonardoi	2+–1–1–0	2–0–0	2–2–0–1+1p+1 2–2–0–1p+1 2–2–0–2	0–2–0+–3p+1	0
Leg III H. italica	1++–1+–1–0	2–0–0	1+–2–2-–2+1p	1+–3–3–3p+1	0–0–1–1
H. fioni	3–2–2–0	2–0–0	2–2–2–1+2p 2–2–2–1p+1+1p 2–2–2–3p	1–3–3–1p+1+2p+1 1–3–3–3p+1	0–0–1–1
H. leonardoi	2+–1+–1–0	2–0–0	2–2–2–2+1p	1–3–3–3p+1	0–0–0+–1
Leg IV H. italica	2+–1–1–0	2–0–0	2–2–2–1p+1+1p 2–2–2–2+1p	1+–3–3–4p+1	0–2–2–2
H. fioni	3–1+–1–0	2–0–0	2–2–2–1+2p 2–2–2–1p+1+2p 2–2–2–3p	2–3–3–1p+1+2p+1	0–1+–2–0++
H. leonardoi	1++–1–1–0	2–0–0	2–2–1+–2+1p	2–3–3–3p+1	0–2–2–2
Discussion