Toxopus politus Bergroth, 1905

Forthman, Michael, Chłond, Dominik & Weirauch, Christiane, 2016, Taxonomic Monograph Of The Endemic Millipede Assassin Bug Fauna Of Madagascar (Hemiptera: Reduviidae: Ectrichodiinae), Bulletin of the American Museum of Natural History 2016 (400), pp. 1-1 : 1-

Toxopus politus Bergroth, 1905 nulate between sternites entirely between II–III

Map 13 and laterally between III–VI. Toxopus signoretii Toxopus politus Bergroth, 1905 , 49: 375. can be further differentiated from T. parkeri by

the sparse vestiture, interocular sulcus near the DISTRIBUTION: According to the original hind margin of the eye, and eye about half width description, the type specimen was collected of the head lengths and not reaching the dorsal from Nosy Be Island off the northwest coast of head surface. The interocular sulcus near the hind Madagascar (map 13). margin of the eye and the eye not reaching the DISCUSSION: We do not redescribe this spe- dorsal but reaching the ventral head surface furcies, since the only known specimen could not ther distinguish T. signoretii from T. basalis . This be located, and none of the specimens examined species differs from T. tibialis by the sparse vestiby us correspond to this species. Many of Ber- ture, ventral medial longitudinal depression on groth’s type specimens are deposited in the Zoo- the head extending onto postocular surface, and logical Museum, University of Helsinki, but eye about half width of the head lengths and not Jansson and Coscarón (1989: appendix I) reaching the dorsal head surface.

reported the female holotype of T. politus to be REDESCRIPTION: MALE: Body length: 12.20 missing (or away on loan) from that collection. mm (holotype). COLORATION: Brownish with The specimen is still absent (Larry Huldén, per- pale yellow postclypeal medial longitudinal sonal commun.). Correspondence with other depression, base of scape, corium basally, tromajor European collections did not offer any fur- chanters, forefemur basally and ventrally, mid ther leads, and we speculate that the type speci- and hind femora except dorsomedially, foretibia men may be lost. The three female specimens of except basally, mid and hind tibiae medially, mid Toxopus examined during our study do not tarsi (fore- and hind missing), anterior half of appear to be conspecific with T. politus based on dorsal laterotergites, and abdominal sternites differences in color patterns, ventral abdominal (except pygophore) paramedially and laterally. structure (segment VI medially longer than V in VESTITURE: Sparse; other features as in generic T. politus ; subequal in female specimens), and description. STRUCTURE: HEAD: Subquadrate locality information. A potential neotype desig- in lateral view (pl. 13K); about as long as wide in nation will therefore have to await availability of dorsal view; ventrally with longitudinal depresadditional Toxopus specimens. sion; anteocular region longer than postocular

(pl. 13K); postclypeus with deep, broad medial Toxopus signoretii (Reuter), 1887 , new longitudinal depression to interocular sulcus;

combination synthlipsis about width of eye; interocular sulcus

Plate 12; map 13 near hind margin of eye (pl. 13B); ocelli large,

located on distinct median tubercle (pl. 13H), Cleptria signoretii Reuter, 1887 , 8: 107.

separated by less than diameter of ocellus; post- REVISED DIAGNOSIS: Males are recognized ocular narrow in dorsal view (pl. 13A); eye about among other Toxopus species by a combination of half of head length, reaching dorsal and ventral the following characters: the coloration, large head surfaces; antenna inserted dorsally on head; body size, subquadrate head shape in lateral view pedicel about as long as scape, curved; labium that is as long as it is wide in dorsal view, eye stout; labial segment III ventrally straight (pl. reaching dorsal head surface, and synthlipsis 13K). THORAX: Anterior pronotal lobe with very about the width of an eye. This species is very small anterolateral projections (pl. 15B); pronotal similar to T. tibialis , T. basalis , and T. parkeri , but transverse furrow divided by paramedian longidiffers by the slightly larger body size, smaller tudinal ridges (pl. 15B); posterior pronotal lobe synthlipsis width, and intersegmental sutures cari- smooth; lateral depressions of posterior pronotal

118 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

lobe smooth; hemelytron nearly reaching dorsal view, and ventrally convex labial segment abdominal apex. ABDOMEN: Apex rounded; III distinguishes this species from T. pallidus . intersegmental sutures carinulate between ster- DESCRIPTION: MALE: Body length: 11.53 mm nites entirely between II–III and laterally (holotype), 10.89–11.62 mm. COLORATION: between III–VI. Blackish with pale yellow antennal segments V DISTRIBUTION: The holotype and one juvenile (except basally) and VI and VII, meso- and were collected from Nosy Be Island off the north- metasterna, coxae, trochanters, forefemur venwest coast of Madagascar (map 13). One speci- trally, mid and hind femora except large subapimen was collected from Vohemar, which is on cal band, mid and hind tibiae medially, tarsi, the northeast coast of Madagascar. anterior one-third of dorsal laterotergites, ventral DISCUSSION: The redescription is based on laterotergites, and abdominal sternites (except habitus images and Reuter’s (1887) original pygophore) medially and laterally. Mandibular description of the holotype. The original descrip- and maxillary plates, postclypeal medial longitution indicates that one immature specimen was dinal depression, labium, posterior pronotal collected (coll. Stumpff) in “Loucoubé,” which is lobe, corium, and lateral longitudinal band currently known as the Natural Reserve of across sternites varying shades of brown. VESTI- Lokobe on Nosy Be Island. Reuter described this TURE: Dense; other features as in generic species in the genus Cleptria Stål, 1859 , but did description. STRUCTURE: HEAD: Ovoid in latnot provide arguments for assigning it to that eral view (pl. 13F); longer than wide in dorsal genus. This species is transferred to Toxopus view; ventrally with deep medial depression; based on the 7-segmented antenna, head being anteocular region longer than postocular (pl. longer than or as long as the width, subequal 13K); postclypeus with deep, narrow medial lonlengths of labial segments II and III, and apter- gitudinal depression to interocular sulcus; synous females, which are features differentiating thlipsis about 1.5 times width of eye; interocular Toxopus from Cleptria . sulcus near hind margin of eye (pl. 13B); post- HOLOTYPE: Male: Madagascar: Antsiranana: ocular narrow in dorsal view (pl. 13A); ocelli Nosy Be, 13.31502 ° S 48.25926 ° E, 155 m (NMW). large, located on distinct median tubercle (pl. PARATYPE: Madagascar: Antsiranana: Nosy Be, 13H), separated by less than diameter of ocellus; 13.31502 ° S 48.25926 ° E, 155 m, 1 juv. (MNHN). eye about one-third of head length, not reaching OTHER MATERIAL EXAMINED: Madagascar:

dorsal and ventral head surfaces; antenna Antsiranana: Vohémar , 13.37028 ° S 50.00337 ° E, 6 m, 13 ( MNHN). Unknown: 13 ( NMW). inserted dorsally on head ; pedicel about one-

fourth longer than scape, slightly curved; labium

stout; labial segment III ventrally convex (pl.

Toxopus simulans , new species 13J). THORAX: Anterior pronotal lobe with small

Plates 4, 7, 11; map 13 anterolateral projections (pl. 15B); posterior pro- DIAGNOSIS: Males recognized among other notal lobe weakly transversely striated; pronotal Toxopus species by the coloration, ovoid head transverse furrow divided by paramedian longishape in lateral view that is longer than wide in tudinal ridges (pl. 15B); lateral depressions of dorsal view, anteocular region longer than post- posterior pronotal lobe weakly transversely striocular, narrow postocular, and labial segment III ated; hemelytron surpassing abdominal apex. ventrally convex. This species is most similar to T. ABDOMEN: Apex rounded; intersegmental pallidus , but the slightly larger body size, ventral sutures carinulate between sternites II and III; medial depression extending along the entire head pygophore process subtriangular in caudal view surface, longer anteocular region relative to the (pl. 19I), not surpassing posterior margin of postocular, interocular sulcus located near the pygophore; BPE as long as basal plate; endohind margin of the eye, narrower postocular in somal struts reaching posterior margin of DPS

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 119

(pl. 21J, K), with anterior and posterior ventral trochanters, base forefemur, forefemur subapiprocesses (pl. 22G); area of endosomal struts– cally and ventrally, mid and hind femora basally DPS fusion short, subquadrate; endosoma medi- and apically, foretibia medially, mid tibia except ally sclerotized (pl. 21L). subbasally and apically, hind tibia basally, fore- ETYMOLOGY: The species epithet is a partici- tarsi, mid and hind tarsi basally, corium basally, ple in the nominative case and is named for the anterior one-third to half of laterotergites, and similar body coloration with Toxopus pallidus . abdominal sternites laterally except posterior DISTRIBUTION: Toamasina and northern two-thirds of segments VI and VII. Maxillary Antsiranana provinces (map 13). plates dorsally, postclypeal longitudinal depres- HOLOTYPE: Male: Madagascar: Antsiranana: sion, paramedially along interocular sulcus, Parc National Montagne d’Ambre, 12.51444 ° S basal half of antennal segment V, labium, mid 49.18138 ° E, 960 m, 04 Mar 2001 – 19 Mar 2001, and hind tarsi apically, and anterior margin of M. Irwin, R. Harin’Hala (00045508) ( CAS). corium (except basally) dark yellow to brown. PARATYPES: Madagascar: Antsiranana: Parc VESTITURE: Dense ; other features as in National Montagne d’Ambre, 12.52027 ° S 49.17916 ° E, generic description. STRUCTURE: HEAD: Sub- 1125 m, 14 May 2001 – 30 May 2001, M. Irwin, R .

quadrate in lateral view (pl. 13K); longer than Harin’Hala, 13 (00007226) ( CAS). Sakalava Beach , dwarf littoral forest, 12.26277 ° S 49.3975 ° E, 10 m, 20 wide in dorsal view ; ventrally with deep longi- Aug 2001 – 28 Aug 2001, M. Irwin, R. Harin’Hala , 13 tudinal depression ; anteocular region longer (00045363) ( CAS). 7 km N of Joffreville , 12.33333 ° S than postocular (pl. 13K) ; postclypeus with 49.25 ° E, 360 m, 13 May 2001 – 16 May 2001, M. Irwin, deep, narrow medial longitudinal depression to

R. Harin’Hala,

13

(00045123) (SU). Toamasina:

interocular sulcus; synthlipsis about two times Botanic Garden near entrance to Andasibe National Park, 18.92633 ° S 48.40783 ° E, 1025 m, 01 Nov 2001 –07 width of eye ; interocular sulcus near hind mar- Nov 2001, M. Irwin, R. Harin’Hala, 13 (00007048) gin of eye (pl. 13B) ; postocular narrow in dorsal (UCR). Unknown: 13 (00045124) (CAS). view (pl. 13A); ocelli large, located on shallow

median tubercle (pl. 13K), separated by less

Toxopus steineri , new species than diameter of ocellus; eye about one-third of

Plates 4, 7, 11; map 12 head length, not reaching dorsal and ventral DIAGNOSIS: Males recognized among other head surfaces; antenna inserted dorsally on species in Toxopus by the coloration, dense ves- head; pedicel about two-fifths longer than titure, narrow postclypeal depression, ventrally scape, slightly curved; labium stout; labial segstraight labial segment III, smooth posterior pro- ment III ventrally convex (pl. 13J). THORAX: notal lobe, and intersegmental sutures carinulate Anterior pronotal lobe with very small anterobetween sternites II–IV and laterally between lateral projections (pl. 15B); posterior pronotal IV–VI. This species is similar to T. namoroka , T. lobe smooth; pronotal transverse furrow basalis , T. vazimba , and T. brucei , but the dark divided by paramedian longitudinal ridges (pl. brown to blackish ventral abdominal surface, 15B); lateral depressions of posterior pronotal narrow postclypeal depression, and ventrally lobe weakly transversely striated; hemelytron straight labial segment III distinguishes T. stein- surpassing abdominal apex. ABDOMEN: Apex eri from these other species. rounded; intersegmental sutures carinulate DESCRIPTION: MALE: Body length: 11.14 between sternites II–IV and laterally between mm (holotype), 11.00– 11.59 mm. COLOR- IV–VI; pygophore process subtriangular in cau- ATION: Blackish with pale yellow bases of dal view (pl. 19I), not surpassing posterior marscape and pedicel, antennal segment V distally gin of pygophore; BPE shorter than basal plate; and segments VI and VII, posterolateral mar- endosomal struts reaching posterior margin of gins of pronotum, lateral areas of transverse DPS (pl. 21J, K), ventrally obscured by semiexsulcus between meso- and metasterna, coxae, tended endosoma; area of endosomal struts–

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DPS fusion subquadrate; endosoma sclerotized num medially, corium basally, mid and hind medially (pl. 21L). coxae, trochanters, base of forefemur, for femur ETYMOLOGY: The species epithet is a noun in ventrally and subbasally, mid and hind femora the genitive case and is named after Ranomafana, except small apical and large medial bands, tibiae Madagascar, which is near the type locality. medially, tarsi, anterior half of laterotergites, and DISTRIBUTION: Known from a single locality abdominal sternites (except pygophore) mediwest of Ranomafana in the Fianarantsoa prov- ally, and lateral margin of sternites. Labium ince (map 12). brown. VESTITURE: Moderately dense ; other HOLOTYPE: Male : Madagascar: Fianarant- features as in generic description. STRUCTURE: soa: 7 km W Ranomafana, 21.25923 ° S HEAD: Subquadrate in lateral view (pl. 13K) ; as 47.38664 ° E, 900 m, 23 Feb 1990 – 28 Feb 1990, long as wide in dorsal view; ventrally with deep W.E. Steiner (00026334) (USNM). anteromedial depression; anteocular region lon- PARATYPES: Madagascar: Fianarantsoa: 7 km W ger than postocular (pl. 13K) ; postclypeus with Ranomafana , 21.25923 ° S 47.38664 ° E, 900 m, 23 Feb deep, broad medial longitudinal depression to 1990– 28 Feb 1990, W.E. Steiner, 33 (00026332,

interocular sulcus; synthlipsis about 1.5 times 00026333, 00026335) (USNM).

width of eye; interocular sulcus near hind mar-

gin of eye (pl. 13B); postocular narrow in dorsal

Toxopus tibialis , new species view (pl. 13A); ocelli large, located on distinct

Plates 4, 7, 11; map 11 median tubercle (pl. 13H), separated by less than DIAGNOSIS: Males are recognized among diameter of ocellus; eye about one-third of head other species in Toxopus by a combination of the length, not reaching dorsal and ventral head surfollowing characters: the relatively larger body faces; antenna inserted dorsally on head; pedicel size, moderately dense vestiture, head as long as about one-fourth longer than scape, slightly it is wide in dorsal view, ventral depression on curved; labium stout; labial segment III ventrally the head restricted to the anteocular region, eye straight (pl. 13K). THORAX: Anterior pronotal about one-third of head length, eye not reaching lobe with very small anterolateral projections (pl. dorsal or ventral head margins, and interseg- 15B); posterior pronotal lobe weakly transversely mental sutures carinulate between sternites II– striated; pronotal transverse furrow divided by VI. Males are most similar to T. basalis and T. paramedian longitudinal ridges (pl. 15B); lateral parkeri , from which it differs by the larger body depressions of posterior pronotal lobe transsize, ventral anteromedial depression on the versely striated; hemelytron reaching or surpasshead, and interocular sulcus located near the ing abdominal apex. ABDOMEN: Apex rounded; hind margin of the eye. The denser vestiture, intersegmental sutures carinulate between stersmaller eye size that does not reach the dorsal nites II–VI; pygophore process subtriangular in and ventral head margins, and intersegmental caudal view (pl. 19I), not surpassing posterior sutures carinulate between sternites II–VI fur- margin of pygophore; BPE as long as basal plate; ther differentiates T. tibialis from T. basalis . A endosomal struts not reaching posterior margin smaller synthlipsis width further distinguished of DPS (pl. 21L), with anterior and posterior T. tibialis from T. parkeri . ventral processes (pl. 22G); area of endosomal DESCRIPTION: MALE: Body length: 14.52 mm struts–DPS fusion elongate ovoid (pl. 21L); (holotype), 13.89–15.40 mm. COLORATION: endosoma medially sclerotized (pl. 21L). Blackish with pale yellow clypeus, dorsal margins ETYMOLOGY: The species epithet is an adjecof mandibular and maxillary plates, postclypeal tive in the nominative case and is named for the medial longitudinal depression, antennifer, scape pale coloration of the tibia. basally, antennal segments V–VII, mesosternum DISTRIBUTION: Antsiranana and Mahajanga except medial longitudinal depression, metaster- provinces in Madagascar (map 11).

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 121

DISCUSSION: The clypeus, mandibular and subbasally and apically, tarsi, anterior half of maxillary plates, and postclypeus are slightly dorsal laterotergites, ventral laterotergites, paradarker in some specimens. medial spots on abdominal sternites II – VI, and HOLOTYPE: Male: Madagascar: Antsiranana: lateral margin of abdomen. Labium, posterolat- Forêt Ambanitaza , 26.1 km 347° Antalaha , eral pronotal lobe, and corium apically brown. 14.67944 ° S 50.18361 ° E, 240 m, 26 Nov 2004, B.L. VESTITURE: Moderately dense ; other features Fisher (00045457) ( CAS). as in generic description. STRUCTURE: HEAD: PARATYPES: Madagascar: Antsiranana: 7 km N of Ovoid in lateral view (pl. 13F) ; as long as wide Joffreville, 12.33333 ° S 49.25 ° E, 360 m, 22 Jan 2001 – 26 in dorsal view; ventrally with deep medial Jan 2001, M.E. Irwin, E.L. Schlinger, R. Harin’Hala , 13

depression; anteocular region longer than post- (00044821) (CAS). Forêt d’Ampondrabe, 26.3 km 10 ° NNE Daraina, 12.97 ° S 49.7 ° E, 175 m, 10 Dec 2003, B.L. ocular (pl. 13K); postclypeus with deep, broad Fisher, 13 (00045701) (CAS). Marojejy National Park, medial longitudinal depression to interocular 5 km W Manantenina village, Camp Mantella, sulcus; synthlipsis about width of eye; interocu- 14.43816 ° S 49.774 ° E, 490 m, 11 Feb 2005 – 18 Feb 2005, lar sulcus near hind margin of eye (pl. 13B); M.

Irwin,

R.

Harin’Hala,

13

(00045492) (SU).

Maha-

postocular narrow in dorsal view (pl. 13A); janga: Réserve forestière Beanka, 50.2 km E Maintirano, 18.02638 ° S 44.05055 ° E, 250 m, 19 Oct 2009 –26 ocelli large, located on distinct median tubercle Oct 2009, B.L. Fisher et al., 13 (00044854) (UCR). (pl. 13H), separated by less than diameter of Unknown: 13 (00045637) (CAS). ocellus; eye about half of head length, reaching

dorsal and ventral head surfaces; antenna

Toxopus toamasina , new species inserted dorsally on head; pedicel about one-

Plates 4, 7, 11; map 10 fourth longer than scape, slightly curved; labium DIAGNOSIS: Males are recognized among other stout ; labial segment III ventrally straight (pl. species in this genus by the coloration, ovoid head 13K). THORAX: Anterior pronotal lobe with shape in lateral view that is as long as it is wide in small anterolateral projections (pl. 15B) ; postedorsal view, synthlipsis about as wide as an eye, rior pronotal lobe transversely striated; pronotal narrow postocular, eye about half the length of the transverse furrow divided by paramedian longihead in lateral view and reaching the dorsal and tudinal ridges (pl. 15B); lateral depressions of ventral head margins, and intersegmental sutures posterior pronotal lobe transversely striated; carinulate between sternites II and III and laterally hemelytron surpassing abdominal apex. ABDObetween III – V. This species is similar to T. simu- MEN: Apex rounded ; intersegmental sutures lans, but differs by the generally darker coloration, carinulate between sternites II and III and laterapex of corium light brown, head as long as it is ally between III–V; pygophore process subtrianwide, smaller synthlipsis width, eye one-half of the gular in caudal view (pl. 19I), not surpassing head length in lateral view, eye height relative to posterior margin of pygophore; BPE as long as dorsal and ventral head margins, labial segment basal plate; endosomal struts reaching posterior III ventrally straight, and intersegmental sutures margin of DPS (pl. 21J, K), with anterior and carinulate between sternites II and III and laterally posterior ventral processes (pl. 22G); area of between III – V. endosomal struts–DPS fusion elongate sub- DESCRIPTION: MALE: Body length: 12.04 quadrate (pl. 21K) ; endosoma medially scleromm (holotype), 11.22–12.41 mm. COLOR- tized (pl. 21L). ATION: Blackish with pale yellow margins of ETYMOLOGY: The species epithet is a noun in ocellar tubercle except posteromedial area, apposition and is named after the Toamasina antennal segments V (except basally) and VI province in Madagascar, where all known speci- and VII, mid and hind coxae, trochanters, fore- mens were collected. femur except dorsally, mid and hind femora DISTRIBUTION: Eastern region the Toamasina except subapically, mid and hind tibiae except province (map 10) .

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DISCUSSION: In some specimens, the scutellar mid tibia subbasally and medially, hind tibia basally processes and abdominal sternite II are pale yel- and subapically, tarsi, anterior half of laterotergites, low, the pale yellow ventrolateral abdominal and abdominal sternites laterally and medially, spots vary in size. except intersternal sutures. Antennal segments V HOLOTYPE: Male: Madagascar: Toamasina: and VI and remainder of corium brown. VESTI- Parcelle E 3, Tampolo, 17.28333 ° S 49.43333 ° E, 10 TURE: Moderately dense ; other features as in m, 14 Apr 2004, Malagasy Ant Team (00006478) generic description. STRUCTURE: HEAD: Cylin- ( CAS). drical in lateral view (pl. 13J) ; longer than wide in PARATYPES: Madagascar: Toamasina: Mobot Site, dorsal view ; ventrally with shallow anteromedial Analalava 7 km SW of Foulpointe , 17.69333 ° S depression ; anteocular region longer than postocu- 49.46027 ° E, 18 m, 03 Jan 2008 – 11 Jan 2008, M. Irwin, R .

lar (pl. 13K); postclypeus with shallow, narrow Harin’Hala, 23 (00007095, 00044942) ( CAS) ; 11 Jan 2008 – 18 Jan 2008, M. Irwin, R. Harin’Hala, 13 medial longitudinal depression to middle of (00006394) ( CAS) ; 22 Feb 2008 – 29 Feb 2008, M. Irwin, interocular area (pl. 13B); synthlipsis about two R. Harin’Hala , 13 (00045546) (SU) ; 04 Apr 2008 –11 Apr times width of eye; interocular sulcus near hind 2008, M. Irwin, R. Harin’Hala , 13 (00007056) (SU) ; 18 margin of eye (pl. 13B); postocular moderately Apr

2008–25

Apr 2008, M.

Irwin,

R.

Harin’Hala,

13

broad in dorsal view; ocelli large, located on shal- (00007078) (UCR); 25 Apr 2008 – 02 May 2008, M. Irwin, R. Harin’Hala, 13 (00045537) ( UCR). Parcelle E 3, Tam- low median tubercle (pl. 13J), separated by less than polo, 17.28333 ° S 49.43333 ° E, 10 m, 14 Apr 2004, Mala- diameter of ocellus ; eye about one-third of head gasy Ant Team, 13 (00007218) ( CAS). length, not reaching dorsal and ventral head sur-

faces; antenna inserted sublaterally on head; pedicel

Toxopus toliara , new species about two-fifths longer than scape, slightly curved;

Plates 4, 7, 11, 21K; map 10 labium slender; labial segment III ventrally straight DIAGNOSIS: Males are recognized among (pl. 13K). THORAX: Anterior pronotal lobe with other Toxopus species by the coloration, moder- small anterolateral projections (pl. 15B); posterior ately dense vestiture, elongate cylindrical head in pronotal lobe smooth; pronotal transverse furrow lateral view, relatively large ocelli, antenna divided by paramedian longitudinal ridges (pl. inserted sublaterally on the head, ventrally 15B); lateral depressions of posterior pronotal lobe straight labial segment III, and smooth posterior weakly to strongly transversely striated; hemelytron pronotal lobe. This species is similar to T. itala- reaching or surpassing abdominal apex. ABDOMEN: viana, T. griswoldi , T. insignis , T. fisheri , and T. Apex rounded; intersegmental sutures carinulate antsiranana , but is distinguished by the diagnos- between sternites II–IV and laterally between IV– tic features mentioned. VI; pygophore process curved in lateral view (pl. DESCRIPTION: MALE: Body length: 11.02 mm 20K), elongate in caudal view (pl. 19H), surpassing (holotype), 10.39–12.54 mm. COLORATION: posterior margin of pygophore; BPE shorter than Blackish with pale yellow maxillary plates dorsally; basal plate; endosomal struts reaching posterior antennifers postclypeal medial longitudinal depres- margin of DPS (pl. 21K), with anterior and postesion, paramedian longitudinal stripes on postclyp- rior ventral processes (pl. 22G); area of endosomal eus anterior of interocular sulcus that extend struts–DPS fusion elongate subquadrate (pl. 21K); laterally along interocular sulcus, labium, scape endosoma with sclerotized, hairlike, denticles on basally, antennal segment VII, collar, pronotum posterior margin (pl. 21K). anterolaterally, anterolateral margin of posterior ETYMOLOGY: The species epithet is a noun in pronotal lobe, meso- and metasterna medially apposition and is named after the Toliara provexcept transverse sulcus, corium basally, coxae, tro- ince in Madagascar, where a majority of the chanters, fore- and mid femora basally and subapi- specimens examined were collected. cally, hind femur except medially, foretibia basally DISTRIBUTION: Fianarantsoa, Mahajanga, and and medially except lateral spots, base of mid tibia, Toliara provinces (map 10).

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 123

DISCUSSION: The coloration is highly variable F. Parker, R. Harin’Hala, 13 (00006615) (CAS); 27 Dec with the pale yellow markings more extensive or 2002– 07 Jan 2003, M. Irwin, F. Parker, R. Harin’Hala, reduced on the head, pronotum, scutellar pro- 33 (00006436, 00006442, 00006446) (USNM), 113

( 00006448, 00006451, 00006601–00006608, 00006664) cesses, and legs. ( CAS); 26 Jan 2003 – 05 Feb 2003, M. Irwin, F. Parker, R. HOLOTYPE: Male: Madagascar: Toliara: Harin’Hala, 73 (00006695–00006701) ( CAS); 24 Mar Berenty Special Reserve , 8 km NW Amboasary, 2003– 03 Apr 2003, M. Irwin, F. Parker, R. Harin’Hala , 25.021 ° S 46.3055 ° E, 35 m, 28 Feb 2004 –24 Mar 63 (00006566–00006570, 00045618) ( CAS); 14 Apr 2004, M. Irwin, F. Parker, R. Harin’Hala 2003– 24 Apr 2003, M. Irwin, F. Parker, R. Harin’Hala,

23 (00005357, 00006624) (AMNH); 24 Apr 2003 –03 (00006598) ( CAS). May 2003, M. Irwin, F. Parker, R. Harin’Hala , 23 PARATYPES: Madagascar: Fianarantsoa: 50 km S of (00006599, 00006600) ( MNHN) ; 14 May 2003 –25 May Farafangana, Mahabo Mananivo, Ampitavananima For- 2003, M. Irwin, F. Parker, R. Harin’Hala , 23 (00006595, est, 23.12983 ° S 47.717 ° E, 34 m, 06 Sep 2007 –13 Sep 00006596) ( CAS) ; 25 May 2003 – 04 Jun 2003, M. Irwin, 2007, M. Irwin, F. Parker, R. Harin’Hala, 13 (00006614) F. Parker, R. Harin’Hala , 13 (00006679) ( UCR) ; 04 Jun ( CAS). Mahajanga: Besalampy District, Marofototra 2003– 10 Jun 2003, M. Irwin, F. Parker, R. Harin’Hala , dry forest, 17 km W of Besalampy, 16.72166 ° S 23 (00006646, 00006678) ( CAS) ; 08 Nov 2003 –15 Nov 44.42366 ° E, 52 m, 19 Nov 2007 – 26 Nov 2007, M. Irwin, 2003, M. Irwin, F. Parker, R. Harin’Hala, 13 (00006677) R. Harin’Hala, 13 (00044907) ( CAS). Toliara: Ando- ( CAS) ; 22 Nov 2003 – 30 Nov 2003, M. Irwin, F. Parker, hahela Natl Park, Tsimelahy, Parcelle II, 24.93683 ° S R. Harin’Hala , 33 (00006525–00006527) ( MNHN) ; 14 46.62666 ° E, 180 m, 27 Dec 2002 – 06 Jan 2003, M. Irwin, Dec 2003 – 21 Dec 2003, M. Irwin, F. Parker, R. F. Parker, R. Harin’Hala , 23 (00006639, 00006643) Harin’Hala, 23 (00006577, 00006578) ( CAS) ; 21 Dec ( CAS) ; 05 Feb 2003 – 15 Feb 2003, M. Irwin, F. Parker, 2003– 02 Jan 2004, M. Irwin, F. Parker, R. Harin’Hala, R. Harin’Hala , 13 (00006674) ( CAS) ; 15 Feb 2003 –26 23 (00006592, 00006593) ( CAS) ; 02 Jan 2004 –13 Jan Feb 2003, M. Irwin, F. Parker, R. Harin’Hala, 13 2004, M. Irwin, F. Parker, R. Harin’Hala , 13 (00006558) (00006628) ( BMNH) ; 18 Mar 2003 – 28 Mar 2003, M. ( SU) ; 13 Jan 2004 – 23 Jan 2004, M. Irwin, F. Parker, R. Irwin, F. Parker, R. Harin’Hala , 13 (00006626) ( CAS) ; Harin’Hala, 23 (00006561, 00006562) ( CAS) ; 04 Feb 29 Jun 2003 – 10 Jul 2003, M. Irwin, F. Parker, R. 2004– 15 Feb 2004, M. Irwin, F. Parker, R. Harin’Hala, Harin’Hala, 63 (00006618, 00006627, 00006629, 23 (00006571, 00006572) ( CAS); 15 Feb 2004 –02 Mar 00006630, 00006632, 00006634) ( USNM) ; 17 Aug 2004, M. Irwin, F. Parker, R. Harin’Hala , 33 (00006619, 2003– 24 Aug 2003, M. Irwin, F. Parker, R. Harin’Hala, 00006621, 00006680) ( BMNH) ; 02 Mar 2004 –11 Mar 13 (00006676) ( CAS) ; 01 Oct 2003 – 11 Oct 2003, M. 2004, M. Irwin, F. Parker, R. Harin’Hala, 23 (00006559, Irwin, F. Parker, R. Harin’Hala , 23 (00006647, 00006560) ( CAS); 05 Apr 2004 – 15 Apr 2004, M. Irwin, 00006669) ( SU) ; 30 Oct 2003 – 09 Nov 2003, M. Irwin, F. Parker, R. Harin’Hala, 23 (00006563, 00006564) F. Parker, R. Harin’Hala , 23 (00006660, 00006666) ( MNHN) ; 30 Apr 2004 – 01 May 2004, M. Irwin, F. ( CAS) ; 30 Nov 2003 – 11 Dec 2003, M. Irwin, F. Parker, Parker, R. Harin’Hala, 13 (00006443) ( SU) ; 14 May R. Harin’Hala, 23 (00006673, 00006675) ( UCR) ; 21 2004– 27 May 2004, M. Irwin, F. Parker, R. Harin’Hala , Dec 2003 – 23 Dec 2003, M. Irwin, F. Parker, R. 43 (00006582–00006585) ( CAS) ; 20 Jun 2004 –30 Jun Harin’Hala, 33 (00006579–00006581) ( AMNH) ; 15 Jan 2004, M. Irwin, F. Parker, R. Harin’Hala , 13 (00006657) 2004– 28 Jan 2004, M. Irwin, F. Parker, R. Harin’Hala, ( CAS) ; 30 Jun 2004 – 11 Jul 2004, M. Irwin, F. Parker, R. 33 (00006609–00006611) ( CAS) ; 28 Jan 2004 –12 Feb Harin’Hala, 23 (00006691, 00006692) ( CAS) ; 25 Jul 2004, M. Irwin, F. Parker, R. Harin’Hala , 63 (00006625, 2004– 07 Aug 2004, M. Irwin, F. Parker, R. Harin’Hala, 00006633, 00006636, 00006638, 00006640, 00006644) 33 (00006574–00006576) ( MNHN) ; 22 Aug 2004 –01 ( BMNH) ; 06 Mar 2004 – 18 Mar 2004, M. Irwin, F. Sep 2004, M. Irwin, F. Parker, R. Harin’Hala, 13 Parker, R. Harin’Hala, 13 (00006617) ( UCR). Berenty (00006565) ( SU) ; 01 Sep 2004 – 08 Sep 2004, M. Irwin, Special Reserve, 8 km NW Amboasary, 25.00666 ° S F. Parker, R. Harin’Hala , 13 (00006591) ( UCR). 46.30333 ° E, 85 m, 25 Oct 2002 – 26 Oct 2002, M. Irwin , Berenty Special Reserve , 8 km NW Amboasary, F. Parker, R. Harin’Hala , 23 (00006649, 00006663) 25.021 ° S 46.3055 ° E, 35 m, 10 Jun 2003 – 19 Jun 2003, M. ( AMNH) ; 02 Nov 2002 – 09 Nov 2002, M. Irwin, F. Irwin, F. Parker, R. Harin’Hala, 13 (00006524) ( CAS) ; Parker, R. Harin’Hala, 23 (00006667, 00006681) ( CAS) ; 22 Nov 2003 – 30 Nov 2003, M. Irwin, F. Parker, R. 30 Nov 2002 – 07 Dec 2002, M. Irwin, F. Parker, R. Harin’Hala, 13 (00006694) ( UCR) ; 07 Dec 2003 –14 Harin’Hala, 13 (00006622) ( UCR) ; 07 Dec 2002 – 14 Dec 2003, M. Irwin, F. Parker, R. Harin’Hala , 23 Dec 2002, M. Irwin, F. Parker, R. Harin’Hala, 13 (00006689, 00006690) ( AMNH) ; 21 Dec 2003 –01 Jan (00006665) ( SU) ; 14 Dec 2002 – 16 Dec 2002, M. Irwin, 2004, M. Irwin, F. Parker, R. Harin’Hala , 23 (00006682,

124 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

00006683) (CAS); 02 Jan 2004 – 13 Jan 2004, M. Irwin, Harin’Hala, 13 (00006661) (CAS); 28 Mar 2002 –08

F. Parker, R. Harin’Hala , 13 (00006686) (SU) ; 15 Feb Apr 2002, M. Irwin, R. Harin’Hala , 23 (00006587,

2004– 28 Feb 2004, M. Irwin, F. Parker, R. Harin’Hala , 00006588) (SU) ; 29 Apr 2002 – 09 May 2002, M. Irwin,

13 (00006620) (CAS); 28 Feb 2004 – 24 Mar 2004, M. R. Harin’Hala, 13 (00006672) (UCR); 06 Mar 2003 –17

Irwin, F. Parker, R. Harin’Hala, 13 (00006597) (CAS); Mar 2003, M. Irwin, R. Harin’Hala, 13 (00006671)

24 Mar 2004 – 04 Apr 2004, M. Irwin, F. Parker, R. (UCR); 27 Jul 2003 – 03 Aug 2003, M. Irwin, R.

Harin’Hala, 23 (00006573, 00006685) (USNM); 04 Apr Harin’Hala, 23 (00006631, 00006655) (CAS).

2004– 15 Apr 2004, M. Irwin, F. Parker, R. Harin’Hala,

13 (00006594) (CAS); 29 Apr 2004 – 11 Jun 2004, M. Toxopus vazimba , new species

Irwin, F. Parker, R. Harin’Hala, 23 (00006612, Plates 4, 7, 11, 19I, 20L, 21L; map 10

00006613) (BMNH); 23 Jun 2004 – 04 Jul 2004, M. DIAGNOSIS: Males recognized among other

Irwin, F. Parker, R. Harin’Hala, 33 (00006468,

00006650, 00006684) (AMNH); 04 Jul 2004 –18 Jul species in Toxopus by the coloration, dense vesti-

2004, M. Irwin, F. Parker, R. Harin’Hala, 13 (00006623) ture, interocular sulcus anterior to the hind mar-

(MNHN); 31 Jul 2004 – 15 Aug 2004, M. Irwin, F. gin of the eye, smooth posterior pronotal lobe,

Parker, R. Harin’Hala, 13 (00006693) (CAS); 29 Aug and intersegmental sutures carinulate between

2004– 08 Sep 2004, M. Irwin, F. Parker, R. Harin’Hala, sternites II–VI and laterally between VI and VII.

13 (00006589) (SU); 08 Sep 2004 – 21 Sep 2004, M. This species is similar to T. steineri , T. namoroka ,

Irwin, F. Parker, R. Harin’Hala, 23 (00006687,

00006688) (CAS); 21 Sep 2004 – 07 Oct 2004, M. Irwin, T. basalis , and T. brucei , but is distinguished from

F. Parker, R. Harin’Hala, 13 (00006590) (UCR). Beza these species by the features mentioned.

Mahafaly Reserve, Parcelle II near Bellevue, 23.68983 ° S DESCRIPTION: MALE: Body length: 11.27 mm

44.5755 ° E, 180 m, 16 Jan 2002 – 18 Jan 2002, M. Irwin, (holotype), 11.27–12.71 mm. COLORATION:

R. Harin’Hala, 13 (00006637) (CAS); 18 Jan 2002 –25 Blackish with pale yellow clypeus, postclypeal

Jan 2002, M. Irwin, R. Harin’Hala, 13 (00044812)

(UCR). Beza Mahafaly Reserve, Parcelle I near research medial longitudinal depression, antennifer, scape

station, 23.6865 ° S 44.591 ° E, 165 m, 15 Oct 2001 –10 basally, pedicel basally, antennal segment VII,

Nov 2001, M.E. Irwin, F.D. Parker, R. Harin’Hala, 13 anterolateral and anteromedial margins of pro-

(00045584) (CAS); 10 Nov 2001 – 21 Nov 2001, R. notum, meso- and metasterna, corium basally,

Harin’Hala, 13 (00006586) (UCR); 21 Nov 2001 –28 coxae, trochanters, forefemur ventrally and

Nov 2001, R. Harin’Hala, 23 (00006099, 00006118) basally, mid and hind femora basally, fore- and

(SU); 28 Nov 2001 – 04 Dec 2001, R. Harin’Hala, 13

(00006653) (CAS); 04 Dec 2001 – 11 Dec 2001, R. mid tibiae except lateral stripes, hind tibia except

Harin’Hala, 23 (00006645, 00006670) (BMNH); 18 apically, tarsi, anterior areas of laterotergites,

Dec 2001 – 25 Dec 2001, R. Harin’Hala, 23 (00006119, abdominal sternites medially, lateral margin of

00006437) (CAS), 23 (00006635, 00006642) (MNHN); sternite II, anterolateral and posterolateral mar-

16 Jan 2002 – 18 Jan 2002, R. Harin’Hala, 23 (00006654, gins of sternites III–VI, and anterolateral margin

00006659) (USNM); 01 Feb 2002 – 08 Feb 2002, R .

Harin’Hala, 13 (00006668) (USNM); 10 Apr 2002 –29 of sternite VII. Labium and antennal segment VI

Apr 2002, R. Harin’Hala, 23 (00006658, 00006662) brown. VESTITURE: Dense; other features as in

(CAS); 08 Jun 2002 – 18 Jun 2002, R. Harin’Hala, 23 generic description. STRUCTURE: HEAD: Sub-

(00006440, 00006444) (AMNH); 28 Jun 2002 –07 Jul quadrate in lateral view (pl. 13K); longer than

2002, R. Harin’Hala, 13 (00006434) (UCR); 07 Jul wide in dorsal view; ventrally with deep medial

2002– 18 Jul 2002, R. Harin’Hala, 13 (00006652) (CAS); depression; anteocular region longer than post-

09 Sep 2002 – 20 Sep 2002, R. Harin’Hala, 23 (00006648,

00006651) (SU); 20 Sep 2002 – 05 Oct 2002, R. ocular (pl. 13K) ; postclypeus with deep, broad

Harin’Hala, 13 (00006656) (CAS); 09 Jan 2003 –23 Jan medial longitudinal depression to interocular

2003, R. Harin’Hala, 13 (00006641) (CAS); 21 Apr sulcus; synthlipsis about two times width of eye;

2003– 29 Apr 2003, R. Harin’Hala, 13 (00006616) interocular sulcus anterior to hind margin of eye;

(UCR). Cap Ste Marie Special Reserve, 74 km S of Tsi- postocular narrow in dorsal view (pl. 13A); ocelli

hombe, 25.58766 ° S 45.163 ° E, 37 m, 02 Jun 2003 –10 Jun

2003, M. Irwin, F. Parker, R. Harin’Hala, 13 (00006447) large, located on shallow median tubercle (pl.

(CAS). Mikea Forest , NW of Manombo, 22.90366 ° S 13K), separated by less than diameter of ocellus ;

43.4755 ° E, 30 m, 06 Jan 2002 – 16 Jan 2002, M. Irwin, R. eye about one-third of head length, not reaching

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 125

dorsal and ventral head surfaces; antenna project. Prior to our monograph, only 10 inserted dorsally on head; pedicel about one- described species of millipede assassin bugs fourth longer than scape, slightly curved; labium were known from the island. Here, we described stout; labial segment III ventrally straight (pl. 63 new species and three new genera of 13K). THORAX: Anterior pronotal lobe with very Ectrichodiinae that are endemic to Madagascar. small anterolateral projections (pl. 15B); poste- This drastic increase in described diversity presrior pronotal lobe smooth; pronotal transverse ents an opportunity to investigate the timing furrow divided by paramedian longitudinal and origin of Madagascan Ectrichodiinae using ridges (pl. 15B); lateral depressions of posterior phylogenetic, divergence dating, and biogeopronotal lobe weakly transversely striated; wings graphic analyses; a companion paper investigatspread out in specimens. ABDOMEN: Apex ing these questions is in review (Forthman and rounded; intersegmental sutures carinulate Weirauch, in review). Phylogenetic studies of between sternites II–VI and laterally between VI Madagascan plants, vertebrates, and some and VII; pygophore process subtriangular in cau- invertebrates support close relationships to dal view (pl. 19I), not surpassing posterior mar- either Afrotropical or Oriental lineages (see gin of pygophore, apex rounded; BPE shorter Vences, 2004; Yoder and Nowak, 2006; Warren than basal plate; endosomal struts not reaching et al., 2010). Assuming their monophyly, Afroposterior margin of DPS (pl. 21L), with anterior tropical-Madagascan relationships are evident and posterior ventral processes (pl. 22G); area of in the genera Glymmatophora and Maraenaspis ; endosomal struts–DPS fusion elongate ovoid (pl. both genera have the bulk of species restricted 21L); endosoma medially sclerotized (pl. 21L). to mainland Africa. Other Madagascan genera, ETYMOLOGY: The species epithet is a noun in e.g., Gibbosella and Marojejycoris , share morapposition and is named after Tombeau Vaz- phological similarities with Afrotropical taxa, imba, Madagascar. e.g., Synavecoris , such as the drab coloration, DISTRIBUTION: Known from Tsingy de Bema- ovoid head shape, and circular spiracles. Howraha National Park in the Mahajanga province in ever, male specimens of Gibbosella also share Madagascar (map 10). some morphological features (e.g., distal part of HOLOTYPE: Male: Madagascar: Mahajanga: M and Cu fused or nearly so; dorsal lateroter- Parc National Tsingy de Bemaraha, 3.4 km 93 ° E gites with posterior tubercles) with Oriental Bekopaka, Tombeau Vazimba, 19.14194 ° S taxa, such as Caecina Stål, 1863 , and Bannania 44.82805 ° E, 50 m, 06 Nov 2001 – 10 Nov 2001, Hsiao, 1973. Only formal phylogenetic analyses Fisheret al. (00006964) (CAS). including comprehensive taxon sampling in the PARATYPES: Madagascar: Mahajanga: Parc National Afrotropical and Oriental regions, as well as Tsingy de Bemaraha, 3.4 km 93 ° E Bekopaka, Tombeau morphological and molecular characters, will Vazimba, 19.14194 ° S 44.82805 ° E, 50 m, 06 Nov 2001 –

shed light on the phylogenetic relationships and 10 Nov 2001, Fisher et al., 13 (00006472) (CAS), 13

(00006730) (SU), 13 (00006967) (UCR). Parc National origins of Madagascan Ectrichodiinae .

Tsingy de Bemaraha, 10.6 km ESE 123° Antsalova,

19.70944 ° S 44.71806 ° E, 150 m, 16 Nov 2001 –20 Nov

ACKNOWLEDGMENTS

2001, Fisheret al., 13 (00007109) (CAS).

We would like to extend our appreciation to the

numerous institutions and colleagues who loaned

CONCLUSION

specimens for this revisionary project. We would The Madagascan invertebrate fauna has not also like to thank curators and staff who assisted us been thoroughly documented, but the situation during the search for type material. We appreciate is improving thanks to large taxonomic surveys the comments on the manuscript and feedback on such as the one that has enabled this taxonomic taxonomic challenges from members of the Weir-

126 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

auch lab, John Heraty, Mark Springer, and Brad Chłond, D. 2011a. Description of two new reduviids Mullens. We thank Doug Yanega for feedback on from Madagascar (Heteroptera: Reduviidae : Stenotaxonomic nomenclature. For financial support, we podainae). Oriental Insects 45: 293–303.

Chłond, D. 2011b. A new species of the Madagascan acknowledge the National Science Foundation

genus Rodepirea Villiers, 1962 ( Hemiptera : Heterop- PEET DEB 093333853 grant (awarded to C.W.) and

tera: Reduviidae : Physoderinae ). Heteropterus funding provided by UCR. Specimens examined Revista de Entomología 11: 235–240.

for this project were acquired under NSF DEB Chłond, D. 2014. A review of the taxonomic position of 0542864 (awarded to Michael Sharkey and Brian Vesbius seyrigi Villiers, 1970 , with a description of Brown) and NSF BSI 0072713 (awarded to Brian two new species of Peyrierocoris Chłond and Junki- Fisher and Charles Griswold). The California Acad- ert, 2010 ( Hemiptera : Heteroptera: Reduviidae : emy of Sciences has graciously donated half of the Harpactorinae). Zootaxa 3785: 288–300.

Chłond, D., and P. Baňař. 2013. Myrmicella , a new paratype material collected by CAS to various

genus of Harpactorinae ( Hemiptera : Heteroptera: American and European collections. We thank Petr

Reduviidae ) from Madagascar. Zootaxa 3718: Baňař and an anonymous reviewer for their valu- 483–495.

able feedback on revisions to this manuscript. Chłond, D., and E. Guilbert. 2012. Sicardicoris , a new

Madagascan subgenus of Peyrierocoris Chłond and

Junkiert 2010 ( Hemiptera : Heteroptera: Reduviidae :

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2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 129

PLATE 1. Dorsal habitus of Gibbosella species , Glymmatophora carolae , and Glymmatophora crassipes .

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PLATE 2. Dorsal habitus of Glymmatophora crassipes , Maraenaspis bidens , Marojejycoris species , and Tanin-