Micrommata Latreille, 1804

Jäger, Peter, 2023, Revision of the huntsman spider genus Micrommata Latreille, 1804 (Sparassidae Sparassinae), Zootaxa 5352 (1), pp. 1-45 : 2-5

Micrommata Latreille, 1804

Type species: Aranea smaragdula Fabricius, 1793 = Micrommata virescens ( Clerck, 1757) (see Jäger 1999a: 6).

Diagnosis. Males ( Figs 4–6 , 19–24 , 41–43 , 56–61 , 84–85 , 104–108 ) are easily distinguished from all other Sparassidae by the spiralled tegulum, the tegular prong and the embolic division with plate, embolus and denticle field; the embolic division is connected to the distal tegulum by a flexible membranous part and is therefore mobile [males of Macrinus Simon, 1887 , Nolavia Kammerer, 2006 and Vindullus Simon, 1880a in the Neotropics also have a spiralled tegulum and a tegular prong, but the long flagelliform embolus is not mobile ( Jäger 2020b: figs 393, 398, 412) and the spiders are not green-coloured or diurnal ( Rheims 2010, 2019; Rheims & Jäger 2008)]. In addition, male genital bulbs lack a conductor [this character state is known only from a few species of the Heteropodinae genus Pseudopoda Jäger, 2000a , that is easily distinguished from Sparassinae , i.e. Micrommata , by the presence of intermarginal denticles and 3 promarginal teeth on the chelicerae ( Jäger 2000a, 2015; Jäger et al. 2015), and most species of the genus Chrosioderma Simon, 1897a and all species of the genus Cebrennus Simon, 1880b ; the latter two distinguished by their brown to yellowish colour ( Silva-Dávila 2005; Jäger 2000c, 2014); and Origes Simon, 1897b , distinguished by the presence of 1 promarginal tooth and 2 retromarginal teeth on chelicerae or the brownish colour pattern ( Jäger & Rheims 2008)]. Females ( Figs 7–12 , 25–32 , 39 , 46–51 , 62–72 , 90–95 , 115–128 ) are distinguished from all other Sparassidae by their roughly heart-shaped, sclerotised median septum (with or without folds), the two distinct pockets at the posterior margin of the epigyne, and the internal duct system with broad membranous parts of the intromittent ducts, a sub-central set of glandular appendages and tight loops, from which long and thin fertilisation ducts emerge, running ventrally from the membranous part in a large semicircle to the epigastric furrow. A superficially similar arrangement can be observed in females of the Neotropical species “ Olios ” erroneus O. Pickard-Cambridge, 1890 , but the two epigynal pockets as well as the glandular appendages or tight loops are missing ( Jäger 2020b: figs 281–286) and the spiders are neither green nor diurnal.

Description. Small to median-sized spiders (TL males 5.1–9.8, females 8.2–17.7). PS longer than wide, fovea distinct, longitudinal, located in posterior half. Head region between 0.5 times (males) and 0.65 times (females) the width of the prosoma. Striae distinct in preserved specimens, not so in live spiders. Prosoma in lateral view with straight outline dorsally until fovea, then steeply sloping almost in a straight line to posterior margin. Eight eyes arranged in two rows, anterior row narrower and recurved, posterior row procurved ( Figs 89 , 100–101 , 110–113 ). AME smallest, ALE largest. Eyes surrounded by white setae (easily rubbed off in preserved material), typical of diurnal spiders. Sternum slightly longer than wide; labium wider than long; gnathocoxae longer than wide. Chelicerae with 2 promarginal, 3–5, rarely up to 7 retromarginal denticles, and 5–10 retromarginal escort setae on the base of the fang. Leg formula: II-IV-I-III or IV-II-I-III. Ta I –IV and MT I–III with sparse scopulae (in MT except for very proximal part); MT IV ventrally with distal patch and a double row of stronger setae, very sparse scopulae present in most distal part. Trilobate membrane with median hook and lateral projections equally strongly developed ( Figs 88 , 96 ). Leg claws with 16–20 secondary teeth, arranged comb-like with primary tooth only slightly longer in males and distinctly longer in females, and female palpal claw with ca. 9 straight teeth and distinctly longer primary tooth ( Figs 86–87 , 97–99 ). OS clearly longer than wide, posterior end slightly pointed.

Palp. The palpal tibia is always shorter than the cymbium and has a simple, pointed, distally arising RTA. The cymbium is characterised by the presence of spines prolaterally and retrolaterally of the alveolus, which are reduced in the majority of Sparassidae . The alveolus is slightly smaller than known from other Sparassidae and is situated relatively centrally within the cymbium. The subtegulum is well visible as a broad ring situated retrolaterally from the tegulum and prolaterally converging in ventral view. The tegulum is twisted with a broad proximal part carrying the broad fundus of the spermophor and a narrowing distal part with a ventral field of denticles terminating in a flat and pointed tegular prong. A conductor is absent. The embolic division is connected to the tegulum by a membranous part near the prong and contains a plate with a second field of denticles and the claw-shaped embolus ( Figs 4–6 , 19–24 , 41–43 , 56–61 , 84–85 , 104–108 ).

Female copulatory organ. The epigynal field is only weakly sclerotised and not always clearly visible. It is roundish and lacks anterior bands. Slit sensilla are more distant from the field than in other Sparassidae and therefore not always observable due to the dissection method. The median septum is strongly sclerotised and has radial grooves in most species and a few transversal grooves in some species. In continuation of the septum there are two epigynal pockets posteriorly, sometimes visible in ventral view, best seen in posterior view. The copulatory openings are slit-like and located laterally to the septum. The IDS begins with a membranous part running from laterally to antero-medially, where it widens into a dorsal pouch, which merges into a narrower sclerotised part with median windings, with a glandular appendage on each side. Fertilisation ducts run from postero-medially first anteriorly, then laterally ventral to the copulatory ducts in a semicircle to the posterior margin ( Figs 7–12 , 25–32 , 39 , 46–51 , 62–72 , 90–95 , 115–128 ).

Reproductive behaviour. The mating and copulatory behaviour of M. virescens was described by Menge (1875), Bristowe (1926) and Nielsen (1932a, b). After detecting a female and a short courtship scene, the male mounts the female in the opposite direction and alternately inserts its genital bulbs into the female’s copulatory opening ( Fig. 135 ). It will direct its body towards the right side of the female’s body and insert its right bulb and its left bulb respectively on the left side of the female’s body. The right bulb is inserted into the right half and the left bulb into the left half of the female’s copulatory organ. µCT scans (Jäger & Michalik, unpublished) show that the male spider first anchors its RTA in one of the epigynal pockets. By expanding the basal haematodocha, the tegular prong is inserted into the slit-like copulatory opening and the entire distal part of the tegulum is inserted into the membranous pouch of the female. This greatly dilates the opening and deforms the median septum. The embolic division is then expanded and the tip of the embolus is in its final position close to the glandular appendages ( Figs 139–140 ).

The whole copulation process can take several hours ( Menge 1875; Jäger, personal observations). During this time, the male may bite the female ( Fig. 136 ), causing injuries, which later become scars ( Figs 134–135 : arrows). In some extreme cases, the female was reported to have died ( Menge 1875). A thorough analysis of all scars found was carried out and will be reported in a separate paper with other species of the spider family Sparassidae (Jäger, in preparation). Most scars were found on one female with 50 scars.

The female will build a retreat by weaving leaves together and will lay eggs inside ( Fig. 137 ). The green eggs are covered within this retreat with only a thin layer of silk ( Fig. 138 ) and are guarded by the female until the spiderlings hatch.

Colouration. Yellowish (preserved material) or green to brown (live spiders). Appendages uniformly coloured, covered with dark setae (spotted in M. aljibica Urones, 2004 ). Dorsal PS with lateral bands of dark spots and dark median stripe or longitudinal patch, the latter continuing onto the dorsal OS in most species (except for M. virescens ). Ventral PS and OS uniformly coloured. It may be that the colour depends on the colour of the vegetation in each latitudinal zone, i.e. the spiders are green where the vegetation is greenish at the peak of the spiders’ life cycle and brown in the more southern parts of the distribution range. Juveniles are coloured differently than adults ( Figs 129–132 ). Body and appendages with variable coverage of dark setae (easily rubbed off in preserved specimens).

Species included. Micrommata aljibica , M. biggi spec. nov., M. diesenhoff spec. nov., M. formosa , M. ligurina , M. virescens .

Notes. “ Micrommata ” darlingi from Zimbabwe is not congeneric with M. virescens and, therefore, not considered here.

Systematic relationships. Member of the family Sparassidae without exact systematic position ( Jäger 1998; Moradmand et al. 2014; Tong et al. 2019; Gorneau et al. 2022). Micrommata was always recovered as separate clade from so-called Sparassinae (i.e. Olios and other genera with 2 promarginal cheliceral teeth, no intermarginal denticles), no sister group could be found so far. If Micrommata proves to be in a clade for its own, this has to be called Sparassinae and the other clade containing above mentioned genera would have to be renamed.

Distribution. Palearctic (without the very northern parts in Europe and Asia), Sierra Leone ( Figs 1–3 , 150 ).

Identification key to Micrommata species

1 Males .............................................................................................. 2

- Females (those specimens of M. aljibica known not fully sclerotised, therefore the respective parts of this key should be treated with caution)......................................................................................... 7

2 Tip of tegular prong retrolaterad in ventral view ( Figs 4 , 57, 59 )................................................ 3

- Tip of tegular prong distad in ventral view ( Figs 20, 23 , 42 , 84 , 105, 107 )......................................... 4

3 Tip of tegular prong long, extending distally beyond alveolus ( Figs 57, 59 ); spiders with distinct dark median band on DS and OS ( Figs 73, 75, 78, 81–83 )........................................................................ formosa

- Tip of tegular prong very short, not extending distally beyond alveolus ( Fig. 4 ); spiders spotted, without distinct median band ( Figs 13–18 ).................................................................................... aljibica

4 Distal margin of tegular prong with incision close to tip in ventral view ( Fig. 84 ); fovea with dark elongate patch ( Fig. 102 )................................................................................................ ligurina

- Distal margin of tegular prong without incision close to tip in ventral view ( Figs 20, 23 , 42 , 105, 107 ); fovea without such patch, in some cases with median band ( Figs 35 , 54 , 135 )........................................................... 5

5 Tip of embolus visible in ventral view, tip of tegular prong roughly as long as the prong’s distal width, tegular loop without or with only few denticles ( Figs 105, 107 ).............................................................. virescens

- Tip of embolus visible in retrolateral view, tip of tegular prong shorter than the prong’s distal width, tegular loop with several to many distinct denticles ( Figs 20–21, 23–24 , 42–43, 44 )..................................................... 6

6 RTA long, i.e. apical tip of RTA reaching tegulum in retrolateral view ( Figs 21, 24 ).............................. biggi

- RTA short, i.e. apical tip of RTA not reaching tegulum in retrolateral view ( Fig. 43 ).......................... diesenhoff

7 Epigynal MS without radial grooves ( Figs 7, 10 , 46 , 48, 51 )................................................... 8

- Epigynal MS with radial grooves ( Figs 25 , 29–31 , 39 , 62, 66 , 68, 71 , 90, 92 )...................................... 9

8 Antero-median ends of epigynal furrows as much separated as lateral lobes at the posterior epigynal margin ( Figs 7, 10 ), IDS with posterior part distinctly bulging laterally and a distinct lateral incision between anterior and posterior part ( Figs 8, 11 )................................................................................................ aljibica

- Antero-median ends of epigynal furrows distinctly closer than lateral lobes at the posterior epigynal margin ( Figs 46 , 48, 51 ), IDS with anterior and posterior part in one longitudinal line, without lateral bulge or incision ( Figs 47–48, 51 ).... diesenhoff

9 Antero-dorsal membranous part of IDS broadly roundish ( Figs 64 , 70 )...................................... formosa

- Antero-dorsal membranous part of IDS tapering or at least not widened ( Figs 27 , 32 , 91, 94 , 116, 118–119 , 123, 126 )..... 10

10 Antero-ventral, narrow part of IDS freely visible in dorsal view ( Figs 116, 118–119 , 123, 126 ).................. virescens

- Antero-ventral, narrow part of IDS covered by dorsal, broad part of IDS in dorsal view ( Figs 27 , 32 , 91, 94 )............ 11

11 Anterior incision of MS V-shaped and with straight and clearly delimited margins, broad dorsal part of copulatory duct with median constriction, anterior ends diverging ( Figs 90, 92 ), fovea marked distinctly with dark longitudinal patch ( Fig. 102 )... ligurina

- Anterior incision of MS indistinctly delimited, broad dorsal part of copulatory duct without median constriction, anterior ends close together ( Figs 25 , 29–31 ), fovea without such patch, but with narrow band with slight waist in front of fovea ( Figs 35, 38 ).............................................................................................. biggi

Figures

FIGURES 4–6. Micrommata aljibica Urones, 2004, holotype male, left palp (4 ventral, 5 retrolateral, 6 tegular tip ventral). EDF—embolic denticle field, EP—embolic plate, ET—embolus tip, F—fundus of spermophore, RTA—retrolateral tibial apophysis, SB—cymbial spine base, SP—spermophor, ST—subtegulum, T—tegulum, TP—tegular prong.

FIGURES 19–24. Micrommata biggi spec. nov., left palp (19, 22 prolateral; 20, 23 ventral; 21, 24 retrolateral) (19–21 holotype male from Ţrkiye; 22–24 male from Turkmenistan). E—embolus, TDF—tegular denticle field, TP—tegular prong.

FIGURES 41–47. Micrommata diesenhoff spec. nov. from Sierra Leone, copulatory organs. 41–45 Left palp (41 prolateral; 42 ventral; 43 retrolateral; 44 tegulum, ventral; 45 embolus tip ventral). 46–47 Female copulatory organ (46 epigyne, ventral; 47 vulva dorsal) (41–45 holotype male; 46–47 female paratype). SO—spermophor opening.

FIGURES 56–61. Micrommata formosa, left palp (56 prolateral; 57, 59 ventral; 58, 60 retrolateral; 61 tegulum tip and embolic division, ventral) (56–58 from Israel; 59–61 male from Spain).

FIGURES 84–89. Micrommata ligurina, male. 84 Left palp, ventral. 85 Ditto, retrolateral. 86–87 leg claws, lateral.88 Trilobate membrane, dorsal. 89 Prosoma with eye arrangement, frontal (84–85 from Spain; 86–89 from Italy: Sardinia). BO—basal outgrowth of embolus, DI—incision of distal tegular margin.

FIGURES 104–109. Micrommata virescens, male. 104–108 Left palp (104 prolateral; 105, 107 ventral; 106, 108 retrolateral). 109 Left chelicera, ventral (104–106 from Russia; 107–108 from Japan; 109 from Germany). E—embolus, EP—embolic plate, TP—tegular prong.

FIGURES 7–12. Micrommata aljibica Urones, 2004, females, copulatory organs (7, 10 epigyne, ventral; 8, 11 vulva dorsal; 9, 12 schematic course of IDS, dorsal) (7–9 paratype, 10–12 MNCN 20.02/19884). adIDS—antero-dorsal part of internal duct system, AE—antero-median ends of epigynal furrows, avIDS—antero-ventral part of internal duct system, CD—copulatory duct, CO—copulatory opening, FD—fertilisation duct, FU—epigynal furrow, GA—glandular appendages, LB—lateral bulge of internal duct system, LL—lateral lobes, MS—median septum; arrows indicating the incision between anterior and posterior part of IDS.

FIGURES 25–28. Micrommata biggi spec. nov., female from Iran, copulatory organ (25 epigyne, ventral; 26 ditto, posterior; 27 vulva dorsal; 28 schematic course of IDS, dorsal). AP—anterior part of internal duct system, EF—epigynal field, EP—epigynal pocket, GA—glandular appendages, MI—antero-median incision of median septum, PMW—postero-median windings, PP— posterior part of internal duct system, RG—radial grooves of median septum.

FIGURES 29–32. Micrommata biggi spec. nov., females from Turkmenistan, copulatory organ (29–31 epigyne, ventral; 32 vulva dorsal).

FIGURES 39–40. Micrommata biggi spec. nov., female paratype from Ţrkiye. 39 Epigyne, ventral. Habitus, frontal.

FIGURES 62–67. Micrommata formosa, female copulatory organ (62, 66 epigyne, ventral; 63 ditto, posterior; 64, 67 vulva, dorsal; 65 schematic course of IDS, dorsal) (62–65 holotype female of M. aragonensis from Spain; 66–67 female from Spain); photo credits: 66–67 Arnaud Henrard. EP—epigynal pocket.

FIGURES 68–72. Micrommata formosa, female copulatory organ (68, 71 epigyne, ventral; 69, 72 ditto, posterior; 70 vulva, dorsal (68–70 from Israel; 71–72 from Tunisia).

FIGURES 90–101. Micrommata ligurina, female. 90–95 Copulatory organ (90, 92 epigyne, ventral; 91, 94 vulva, dorsal; 93 epigyne, posterior; 95 schematic course of IDS, dorsal). 96 Trilobate membrane, dorsal. 97–98 Leg claws, lateral. 99 Palpal claw, lateral. 100–101 Prosoma with eye arrangement (100 frontal; 101 lateral) (90–95 from Spain; 96–101 from Italy: Sardinia).

FIGURES 115–120. Micrommata virescens, female copulatory organ (115, 117 epigyne, ventral; 116, 118–119 vulva, dorsal; 120 schematic course of IDS, dorsal) (115–116 from Russia; 117–120 from Japan).

FIGURES 121–128. Micrommata virescens, female copulatory organ (121, 125 127–128 epigyne, ventral; 122 ditto, posterior; 123, 126 vulva, dorsal; 124 schematic course of IDS, dorsal) (121–124 from Germany; 125–127 from Kazakhstan; 128 from Russia).

FIGURES 110–114. Micrommata virescens from Germany. 110–113 Prosoma with eye arrangement (110, 113 dorsal; 111 lateral; 112 frontal). 114 Left chelicera, ventral (110–112 male; 113–114 female).

FIGURES 133–138. Micrommata virescens, adults from Germany, habitus and behaviour. 133 Male, looking for a female. 134 Female and male in the natural habitat. 135 Male and female in mating position. 136 Ditto, male bites leg of female. 137 Female with egg sac in rolled leaf of Rubus sp. 138 Egg sac with green eggs. Arrows indicating scars on the opisthosoma of female spiders. Photo credit: 136 Herbert Schirmer.

FIGURES 139–140. Micrommata virescens, µ-CT pictures of male and female genitalia during copulation (139 dorsal; 140 lateral). Pink: female internal duct system; light blue: RTA; light violet: cymbium plus basal haematodocha; dark blue: subtegulum and tegulum; black: tegular prong; golden: embolic division.

FIGURES 129–132. Micrommata virescens, immatures from Germany, habitus (129, 132 juvenile; 130–131 subadult male).

FIGURES 1–3. Distribution ranges of Micrommata species. 1 M. virescens (red circles), M. diesenhoff spec. nov. (black triangle). 2 M.ligurina (red circles), M. biggi spec. nov. (black triangles).3 M. formosa (red circles), M.aljibica (black triangles). Majority of records is from specimens examined, some are from published literature or websites like iNaturalist (western record of M. aljibica, several records of M. virescens in its central range).

FIGURES 149–150. 149 Distribution records of Sparassidae across the world excluding the genus Micrommata. The map is based on almost 4300 data sets from literature and spiders examined and does not include all taxa. 150 Elevational distribution of Micrommata spiders from the literature and examined in this study.

FIGURES 73–83. Micrommata formosa, habitus of preserved (73–79) and live (80–83) spiders (73, 75, 78, 80, 82–83 dorsal; 74, 76, 79 ventral; 77 lateral; 81 frontal) (73–74 male from Spain; 75–77 holotype female of M. aragonensis from Spain; 78–81 female from Spain; 82 female from Malta; 83 female with opened egg sac from Portugal). Photo credits: 78–81 Arnaud Henrard; 82 Victor Falzon; 83 Sergio Henriques.

FIGURES 13–18. Micrommata aljibica Urones, 2006, habitus. 13 Holotype male, dorsal. 14–15 Female from NW of Almeria, Spain (14 dorsal, 15 frontal). 16–18 Female from ca. 10 km SE of the type locality (16–17 dorsal, 18 ventral). Photo credits: 14–15 Paco Faluke; 16–18 Oscar Diaz.

FIGURES 102–103. Micrommata ligurina, male from Greece (Corfu), live spider, dorsal. Arrow indicating dark foveal patch.

FIGURES 33–38. Micrommata biggi spec. nov., habitus of preserved specimens from Ţrkiye (33, 35–36, 38 dorsal; 34, 37 ventral) (33–35 holotype male; 36–38 female paratype). Arrows indicating slight waist of median band in front of fovea.

FIGURES 54–55. Micrommata diesenhoff spec. nov., habitus of preserved specimens from Sierra Leone, dorsal (54 holotype male; 55 female paratype).

FIGURES 48–53. Micrommata diesenhoff spec. nov., female paratypes from Sierra Leone, copulatory organs (48, 51 epigyne, ventral; 49, 53 vulva dorsal; 50 schematic course of IDS, dorsal; 52 epigyne, posterior). SS—slit sensillum.