Tanindrazanus marginatus, Forthman & Chłond & Weirauch, 2016

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-

Tanindrazanus marginatus , new species lar in lateral view (pl. 20H, I); BPE longer than

Plates 3, 6, 9; map 8 basal plate; area of endosomal struts–DPS fusion DIAGNOSIS: Males are recognized among ovate (pl. 21H). other species in this genus by the brownish col- ETYMOLOGY: The species epithet is an adjecoration, synthlipsis about the width of an eye, tive in the nominative case and is named for the and large eyes reaching the dorsal and ventral pale laterotergites. head surfaces. This species is similar to T. tene- DISTRIBUTION: Antsiranana, Fianarantsoa, bricus and T. nigripes , but differs from both spe- Toamasina, and Toliara provinces (map 8). cies by the lighter coloration and lateral DISCUSSION: In some specimens, the dorsal carinulation on the intersegmental suture neck surface, thorax, corium, coxae, and laterobetween sternites VI and VII. Tanindrazanus tergites are light brown and the postclypeus marginatus is further distinguished from T. medially dark brown. nigripes by the smaller synthlipsis width. HOLOTYPE: Male: Madagascar: Fianarant- DESCRIPTION: Medium body size (length: soa: Parc National Ranomafana, Vohiparara, at 16.43 mm, holotype; 13.62–17.43 mm). COLOR- broken bridge, 21.22616 ° S 47.36983 ° E, 1110 m, ATION: Dark brown with light brown clypeus, 08 Apr 2002 – 15 Apr 2002, M. Irwin, R. postclypeus medially, dorsal margin of ocellar Harin’Hala (00006821) (CAS). tubercle, labium, basal wing veins, trochanters, PARATYPES: Madagascar: Antsiranana: Marojejy femora except apically, tibiae basally, tarsi, and National Park, 5 km W Manantenina village, Camp

Mantella, 14.43816 ° S 49.774 ° E, 490 m, 04 Feb 2005 –11 lateral margin of sternites. Distal half of antennal

Feb 2005, M. Irwin, R. Harin’Hala, 13 (00006892) segment V and segments VI and VII white. VES- (CAS); 18 Mar 2005 – 25 Mar 2005, M. Irwin, R. TITURE: As in generic description. STRUC- Harin’Hala, 13 (00006888) (CAS); 28 Apr 2005 –07 TURE: HEAD: Ventrally with shallow May 2005, M. Irwin, R. Harin’Hala, 13 (00006908) anteromedial depression; clypeal apex dorsally (USNM); 13 Jul 2005 – 28 Jul 2005, M. Irwin, R. elevated relative to labrum (pl. 13G, H); postclyp- Harin’Hala, 13 (00006087) (AMNH); 14 Oct 2005 –22

Oct 2005, M. Irwin, R. Harin’Hala, 13 (00006850) eus with relatively deep, broad medial longitudi- (SU). Montaigne Francais, 12.325 ° S 49.33333 ° E, 150 m, nal depression to middle of interocular area; 30 Jan 2001 – 15 Feb 2001, M. Irwin, R. Harin’Hala, 23 synthlipsis about width of eye; interocular sulcus (00006833, 00006847) (CAS), 13 (00006921) (AMNH); near hind margin of eye (pl. 13B); postocular 06 Mar 2001 – 20 Mar 2001, M. Irwin, R. Harin’Hala, narrow in dorsal view (pl. 13A); ocelli large, sep- 13 (00006920) (UCR). Parc National Montagne

d’Ambre, 12.51444 ° S 49.18138 ° E, 960 m, 12 Feb 2001 – arated by less than diameter of ocellus, located

04 Mar 2001, M. Irwin, R. Harin’Hala, 13 (00006932) on distinct median tubercle (pl. 13H); eye about (CAS). Parc National Montagne d’Ambre, 12.52027 ° S one-third of head length, reaching dorsal and 49.17916 ° E, 1125 m, 30 May 2001 – 06 Jun 2001, M. ventral head surfaces; pedicel about one-fourth Irwin, R. Harin’Hala, 13 (00006834) (UCR). Fianaralonger than scape; labial segment III subequal to ntsoa: 50 km S of Farafangana, Mahabo Mananivo, II. THORAX: Anterior pronotal lobe with very Ampitavananima Forest, 23.12983 ° S 47.717 ° E, 34 m, 13

Jan 2007– 20 Jan 2007, M. Irwin, F. Parker, R. Harin’Hala, small anterolateral projections (pl. 15B); poste- 13 (00006931) (MNHN); 04 Feb 2007 – 10 Feb 2007, M. rior pronotal lobe transversely striated; pronotal Irwin, F. Parker, R. Harin’Hala, 43 (00006805, transverse furrow continuous; legs slender; tar- 00006806, 00006855, 00006860) (CAS); 10 Feb 2007 –17 someres I and II combined subequal to III on Feb 2007, M. Irwin, F. Parker, R. Harin’Hala, 33 forelegs, slightly longer on mid and hind legs; (00006803, 00006804, 00006861) (AMNH); 03 Mar

2007– 10 Mar 2007, M. Irwin, F. Parker, R. Harin’Hala, hemelytron reaching or surpassing abdominal

13 (00006941) (UCR); 17 Mar 2007 – 24 Mar 2007, M. apex. ABDOMEN: Apex rounded; dorsal lateroter- Irwin, F. Parker, R. Harin’Hala, 13 (00006936) (SU); 24 gite II not expanded; intersegmental sutures cari- Mar 2007– 31 Mar 2007, M. Irwin, F. Parker, R. nulate between sternite II–IV and laterally Harin’Hala, 13 (00006810) (CAS); 07 Apr 2007 –14 Apr between IV–VII; pygophore process subtriangu- 2007, M. Irwin, F. Parker, R. Harin’Hala, 13 (00006937)

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 83

(CAS); 14 Apr 2007 – 21 Apr 2007, M. Irwin, F. Parker, (00045415) ( CAS) ; 28 Jan 2002 – 04 Feb 2002, M. Irwin, R. Harin’Hala, 13 (00006942) ( CAS) ; 06 May 2007 –14 R. Harin’Hala, 23 (00006879, 00006880) ( CAS) ; 12 Feb May 2007, M. Irwin, F. Parker, R. Harin’Hala, 13 2002– 19 Feb 2002, M. Irwin, R. Harin’Hala , 23 (00006801) ( BMNH) ; 14 May 2007 – 21 May 2007, M. (00006835, 00006940) (SU) ; 14 Apr 2002 – 23 Apr 2002, Irwin, F. Parker, R. Harin’Hala, 13 (00006943) M. Irwin, R. Harin’Hala , 13 (00045707) ( CAS) ; 03 Jun ( MNHN) ; 02 Jun 2007 – 10 Jun 2007, M. Irwin, F. 2002– 13 Jun 2002, M. Irwin, R. Harin’Hala, 13 Parker, R. Harin’Hala , 13 (00006930) ( CAS) ; 07 Jul (00006904) ( MNHN) ; 13 Jun 2002 – 23 Jun 2002, M. 2007– 14 Jul 2007, M. Irwin, F. Parker, R. Harin’Hala, Irwin, R. Harin’Hala , 13 (00006905) ( CAS) ; 16 Feb 13 (00006934) ( CAS) ; 14 Jul 2007 – 19 Jul 2007, M. 2003– 26 Feb 2003, M. Irwin, R. Harin’Hala, 23 Irwin, F. Parker, R. Harin’Hala , 13 (00006926) (SU) ; 19 (00006883, 00006884) (USNM); 21 Mar 2003 –12 Apr Jul 2007– 23 Jul 2007, M. Irwin, F. Parker, R. Harin’Hala, 2003, M. Irwin, R. Harin’Hala , 13 (00006891) 13 (00006927) (SU) ; 23 Jul 2007 – 28 Jul 2007, M. ( BMNH) ; 04 May 2003 – 16 May 2003, M. Irwin, R. Irwin, F. Parker, R. Harin’Hala , 13 (00006928) Harin’Hala, 23 (00006885, 00006886) ( AMNH) ; 15 (AMNH); 03 Aug 2007 – 11 Aug 2007, M. Irwin, F. May 2003– 28 May 2003, M. Irwin, R. Harin’Hala, 13 Parker, R. Harin’Hala , 43 (00006811, 00006857, (00006960) ( CAS); 28 May 2003 – 06 Jun 2003, M. 00006858, 00006938) ( USNM) ; 27 Aug 2007 –06 Sep Irwin, R. Harin’Hala, 13 (00045034) ( UCR). Parc 2007, M. Irwin, F. Parker, R. Harin’Hala , 13 (00006929) National Ranomafana, Vohiparara, at broken bridge, ( CAS) ; 06 Sep 2007 – 13 Sep 2007, M. Irwin, F. Parker, 21.22616 ° S 47.36983 ° E, 1110 m, 19 Feb 2002 –26 Feb R. Harin’Hala, 23 (00006856, 00006944) ( MNHN) ; 27 2002, M. Irwin, R. Harin’Hala, 13 (00006902) ( CAS) ; Sep 2007– 04 Oct 2007, M. Irwin, F. Parker, R. 04 Mar 2002 – 12 Mar 2002, M. Irwin, R. Harin’Hala , Harin’Hala, 33 (00006807–00006809) ( CAS) ; 25 Oct 13 (00006837) (SU); 26 Mar 2002 – 31 Mar 2002, M. 2007– 08 Nov 2007, M. Irwin, F. Parker, R. Harin’Hala, Irwin , R. Harin’Hala, 33 (00006844–00006846) ( CAS), 13 (00006922) ( CAS) ; 08 Nov 2007 – 22 Nov 2007, M. 13 (00006924) ( USNM) ; 31 Mar 2002 – 08 Apr 2002, M. Irwin, F. Parker, R. Harin’Hala, 33 (00006812, Irwin, R. Harin’Hala, 13 (00006903) ( USNM); 08 Apr 00006859, 00006945) ( BMNH) ; 28 Nov 2007 – 13 Dec 2002 – 15 Apr 2002, M. Irwin, R. Harin’Hala, 13 2007, M. Irwin, F. Parker, R. Harin’Hala, 13 (00006935) (00006799) ( AMNH), 173 (00006815–00006820, ( USNM); 03 Jan 2008 – 09 Jan 2008, M. Irwin , F. Parker, 00006822–00006830) ( CAS), 33 (00006843, 00006925, R. Harin’Hala, 1♂ (00006907) (SU); 09 Jan 2008 –17 Jan 00006933) ( UCR). Parc National Ranomafana , radio 2008, M. Irwin , F. Parker, R. Harin’Hala , 13 (00006836) tower at forest edge, 21.251 ° S 47.40716 ° E, 1130 m, 06 ( UCR) ; 01 Apr 2008 – 09 Apr 2008, M. Irwin, F. Parker, Dec 2001– 15 Dec 2001, M. Irwin, R. Harin’Hala, 13 R. Harin’Hala, 23 (00006887, 00006890) ( CAS). (00006917) ( CAS) ; 21 Dec 2001 – 24 Dec 2001, M. Manombo Special Reserve camp site, 32 km SSE of Irwin , R. Harin’Hala, 13 (00044908) ( UCR) ; 14 Jan Farafangana, 23.02183 ° S 47.72 ° E, 36 m, 10 Oct 2004 –21 2002– 21 Jan 2002, M. Irwin, R. Harin’Hala, 13 Oct 2004, M. Irwin, R. Harin’Hala , 13 (00045347) (00045718) ( CAS) ; 28 Jan 2002 – 04 Feb 2002, M. Irwin , ( CAS) ; 23 Nov 2004 – 05 Dec 2004, M. Irwin, R. R. Harin’Hala , 13 (00099047) ( CAS) ; 09 Apr 2002 –16 Harin’Hala, 13 (00045266) ( UCR) ; 15 Dec 2004 – 26 Apr 2002, M. Irwin, R. Harin’Hala, 13 (00006896) Dec 2004, M. Irwin, R. Harin’Hala , 13 (00045207) ( USNM) ; 30 Apr 2002 – 07 May 2002, M. Irwin , R. ( AMNH) ; 26 Dec 2004 – 05 Jan 2005, M. Irwin, R. Harin’Hala, 13 (00006839) ( AMNH) ; 07 May 2002 –14 Harin’Hala, 33 (00045006, 00045195, 00045522) May 2002, M. Irwin, R. Harin’Hala, 13 (00006802) ( CAS) ; 16 Jan 2005 – 20 Jan 2005, M. Irwin , R. ( CAS) ; 18 Feb 2003 – 27 Feb 2003, M. Irwin, R. Harin’Hala , 13 (00045467) (SU) ; 15 Feb 2005 –27 Feb Harin’Hala, 13 (00007112) ( MNHN) ; 09 Mar 2003 –20 2005, M. Irwin, R. Harin’Hala, 23 (00044964, Mar 2003, M. Irwin, R. Harin’Hala , 13 (00006895) 00045376) ( CAS) ; 24 Apr 2005 – 10 May 2005, M. Irwin , ( BMNH) ; 07 May 2003 – 17 May 2003, M. Irwin, R. R. Harin’Hala , 13 (00044810) ( BMNH) ; 10 May 2005 – Harin’Hala, 23 (00006874, 00006875) (SU) ; 06 Aug 22 May 2005, M. Irwin, R. Harin’Hala, 13 (00045152) 2003– 16 Aug 2003, M. Irwin, R. Harin’Hala , 33 ( BMNH) ; 06 Jun 2005 – 24 Jun 2005, M. Irwin , R. (00006911–00006913) ( CAS), 13 (00006919) ( BMNH) ; Harin’Hala, 23 (00007190, 00045382) ( USNM) ; 24 Jun 28 Sep 2003 – 08 Oct 2003, M. Irwin, R. Harin’Hala, 13 2005– 10 Jul 2005, M. Irwin, R. Harin’Hala , 13 (00006898) ( CAS) ; 18 Oct 2003 – 26 Oct 2003, M. Irwin , (00044842) ( MNHN) ; 10 Jul 2005 – 27 Jul 2005, M. R. Harin’Hala, 13 (00006899) ( CAS) ; 17 Dec 2003 –30 Irwin , R. Harin’Hala, 13 (00045119) ( CAS) ; 09 Oct Dec 2003, M. Irwin, R. Harin’Hala, 23 (00006831, 2005– 16 Oct 2005, M. Irwin, R. Harin’Hala , 13 00006832) ( CAS) ; 30 Dec 2003 – 11 Jan 2004, M. Irwin, (00044946) ( UCR). Parc National Ranomafana, Belle R. Harin’Hala , 13 (00006914) ( CAS) ; 11 Jan 2004 –21 Vue at Talatakely, 21.2665 ° S 47.42016 ° E, 1020 m, 15 Jan 2004, M. Irwin, R. Harin’Hala, 23 (00006881, Nov 2001– 22 Nov 2001, M. Irwin, R. Harin’Hala , 13 00006882) ( MNHN) ; 31 Jan 2004 – 12 Feb 2004, M.

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Irwin, R. Harin’Hala, 13 (00006894) (CAS); 21 Mar 00006909) (CAS), 13 (00044801) (USNM); 25 Jan

2004– 02 Apr 2004, M. Irwin, R. Harin’Hala, 23 2008– 01 Feb 2008, M. Irwin, R. Harin’Hala, 13

(00006796, 00006797) (AMNH); 02 Apr 2004 –15 Apr (00006923) (CAS), 13 (00006939) (AMNH); 21 Mar

2004, M. Irwin, R. Harin’Hala, 13 (00006840) (USNM); 2008– 28 Mar 2008, M. Irwin, R. Harin’Hala, 13

24 Jun 2004 – 08 Jul 2004, M. Irwin, R. Harin’Hala, 23 (00006798) (UCR). Mobot Site, Analalava 7 km SW of

(00006838, 00006897) (BMNH); 07 Oct 2004 –20 Oct Foulpointe, 17.70889 ° S 49.45806 ° E, 24 m, 15 Sep 2007 –

2004, M. Irwin, R. Harin’Hala, 13 (00006918) (UCR); 21 Sep 2007, M. Irwin, R. Harin’Hala, 13 (00006906)

14 Jan 2005 – 02 Feb 2005, M. Irwin, R. Harin’Hala, 13 (CAS). Toliara: Kirindy forest, 60 km NE of Moron-

(00006868) (SU); 27 Feb 2005 – 13 Mar 2005, M. Irwin, dava, 20.0665 ° S 44.65767 ° E, 45 m, 18 Oct 2003 –30 Oct

R. Harin’Hala, 33 (00006841, 00006842, 00006916) 2003, M. Irwin, R. Harin’Hala, 13 (00045067) (CAS).

(CAS); 28 Mar 2005 – 10 Apr 2005, M. Irwin, R. Unknown: 33 (00006813, 00006814, 00006849) (CAS).

Harin’Hala, 13 (00006900) (BMNH); 25 Sep 2005 –07

Oct 2005, M. Irwin, R. Harin’Hala, 13 (00006869)

(CAS); 06 Nov 2005 – 13 Nov 2005, M. Irwin, R. Tanindrazanus marojejy , new species

Harin’Hala, 13 (00006946) (CAS); 13 Nov 2005 –24 Plates 3, 6, 9, 20H; map 6

Nov 2005, M. Irwin, R. Harin’Hala, 13 (00006865) DIAGNOSIS: Males recognized among other

(MNHN); 21 Dec 2005 – 30 Dec 2005, M. Irwin, R. Tanindrazanus species by the black body with

Harin’Hala, 13 (00006866) (BMNH); 20 Jan 2006 –29 red abdomen and pale orange hemelytra, ven-

Jan 2006, M. Irwin, R. Harin’Hala, 23 (00006872,

00006873) (SU); 29 Jan 2006 – 07 Feb 2006, M. Irwin, R. trally flat head surface, broad postocular in dor-

Harin’Hala, 13 (00006915) (CAS); 15 Feb 2006 –26 Feb sal view, labial segment II shorter than segment

2006, M. Irwin, R. Harin’Hala, 93 (00006947– III, and continuous pronotal transverse furrow.

00006950, 00006953–00006957) (CAS); 05 Mar 2006 – This species is similar to T. anjozorobeus and T.

18 Mar 2006, M. Irwin, R. Harin’Hala, 23 (00006854, kathrynae , but is distinguished by the black head,

00006867) (UCR); 18 Mar 2006 – 30 Mar 2006, M.

Irwin, R. Harin’Hala, 33 (00006862–00006864) thorax, and legs; pale orange hemelytra; flat ven-

(MNHN); 13 Apr 2006 – 24 Apr 2006, M. Irwin, R. tral head surface; synthlipsis about 1.5 times

Harin’Hala, 13 (00006870) (AMNH); 02 May 2006 –13 width of eye; labial segment II shorter than III;

May 2006, M. Irwin, R. Harin’Hala, 23 (00006871, and intersegmental sutures carinulate between II

00006901) (MNHN); 20 May 2006 – 29 May 2006, M. and III and laterally between III–VI.

Irwin, R. Harin’Hala, 13 (00006893) (CAS); 09 Aug DESCRIPTION: Medium body size (length:

2006– 23 Aug 2006, M. Irwin, R. Harin’Hala, 13

(00006800) (USNM); 23 Aug 2006 – 07 Sep 2006, M. 17.53 mm, holotype; 17.05–19.72 mm). COLOR-

Irwin, R. Harin’Hala, 13 (00006853) (CAS); 07 Sep ATION: Black with antennal segment VI and

2006– 20 Sep 2006, M. Irwin, R. Harin’Hala, 33 basal half of segment VII, hemelytron, tarsi, and

(00006851, 00006852, 00006951) (BMNH); 20 Sep pretarsi orange and distal half of antennal seg-

2006– 05 Oct 2006, M. Irwin, R. Harin’Hala, 13 ment VII pale orange. Abdomen red with ante-

(00006952) (SU); 01 Nov 2006 – 11 Nov 2006, M. Irwin,

R. Harin’Hala, 13 (00044923) (UCR). Parc National rior margin of sternite II, sternite VII (except

d’Isalo, 9.1 km 354° N Ranohira, 22.48166 ° S 45.46166 ° E, small anterolateral spot), laterotergite VII (except

725 m, 27 Jan 2003 – 31 Jan 2003, Fisher et al., 13 anterior margin), and pygophore black. VESTI-

(00005363) (AMNH), 33 (00006876–00006878) TURE: As in generic description. STRUCTURE:

(CAS). Ranomafana JIRAMA water works, 21.2485 ° S HEAD: Ventrally flat ; clypeal apex dorsally ele-

47.45216 ° E, 690 m, 16 Oct 2001 – 08 Nov 2001, M. vated relative to labrum (pl. 13G, H); synthlipsis

Irwin, R. Harin’Hala, 13 (00005361) (CAS). Toama-

sina: Botanic Garden near entrance to Andasibe about 1.5 times width of eye; interocular sulcus

National Park, 18.92633 ° S 48.40783 ° E, 1025 m, 01 Sep posterior to hind margin of eye; ocelli large, sep-

2001– 05 Sep 2001, M. Irwin, F. Parker, R. Harin’Hala, arated by less than diameter of ocellus, located

13 (00006889) (CAS). Fampanambo, 15.37550 ° S on distinct median tubercle (pl. 13H); postclyp-

49.62175 ° E, 108 m, 1962, J. Vadon, 13 (MRAC). Mobot eus with shallow, broad depression to middle of

Site, Analalava 7 km SW of Foulpointe, 17.69333 ° S

49.46027 ° E, 18 m, 21 Dec 2007 – 28 Dec 2007, M. Irwin, interocular area (pl. 13B); postocular moderately

R. Harin’Hala, 13 (00044874) (SU); 18 Jan 2008 –25 Jan broad in dorsal view; eye about one-fourth of

2008, M. Irwin, R. Harin’Hala, 23 (00006848, head length, not reaching dorsal and ventral

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 85

head surfaces; pedicel about one-fourth longer ocellar tubercle, and fore- and mid femora venthan scape; labial segment III longer than II. trally and laterally pale yellow. Labium light THORAX: Anterior pronotal lobe without distinct brown except segment II basally and distal twoanterolateral projections; posterior pronotal lobe thirds of antennal segment V and segments VI transversely striated; pronotal transverse furrow and VII white. VESTITURE: As in generic continuous ; legs slender; tarsomeres I and II description. STRUCTURE: HEAD: Ventrally combined subequal to III; hemelytron almost with shallow anteromedial depression; clypeal reaching or surpassing abdominal apex. ABDO- apex dorsally elevated relative to labrum (pl. MEN: Apex rounded; dorsal laterotergite II not 13G, H); postclypeus with shallow, broad medial expanded; intersegmental sutures carinulate longitudinal depression to middle of interocular between II and III and laterally between III–VI; area (pl. 13B); synthlipsis about 1.5 times width pygophore process subtriangular in lateral view of eye; interocular sulcus at (pl. 13B) or poste- (pl. 20H); BPE shorter than basal plate; area of rior to hind margin of eye; postocular narrow in endosomal struts–DPS fusion elongate subquad- dorsal view (pl. 13A); ocelli large, separated by rate (pl. 21G). less than diameter of ocellus, located on distinct ETYMOLOGY: The species epithet is a noun in median tubercle (pl. 13H) ; eye about one-third apposition and is named after the type locality, of head length, not reaching dorsal and ventral Marojejy Nature Reserve , Madagascar. head surfaces ; pedicel about two-fifths longer DISTRIBUTION: Marojejy Nature Reserve in than scape ; labial segment III subequal to II. the Antsiranana province (map 6). THORAX: Anterior pronotal lobe with very small HOLOTYPE: Male : Madagascar: Antsiranana: anterolateral projections (pl. 15B) ; posterior R. N.I. de Marojejy, 11.0 km NW Manantenina , pronotal lobe transversely striated ; pronotal 14.43667 ° S 49.74167 ° E, 1225 m, 25 Oct 1996 –03 transverse furrow divided by paramedian longi- Nov 1996, Eric L. Quinter (00078359) ( AMNH). tudinal ridges (pl. 15B) ; forefemur moderately PARATYPES: Madagascar: Antsiranana: Sambava incrassate ; tarsomeres I and II combined sub- District, R.N. XII Marojejy, Ambatosoratra, equal to III on fore- and mid legs, longer on 14.38735 ° S 49.75717 ° E, 1700m, Nov 1960, P. Soga ,

hind legs; hemelytron surpassing abdominal 33 (MNHN).

apex. ABDOMEN: Apex rounded; dorsal latero-

tergite II not expanded; intersegmental sutures

Tanindrazanus nigripes , new species carinulate between sternites II–IV; pygophore

Plates 3, 6, 9, 22E; map 8 process subtriangular in lateral view (pl. 20H, I); DIAGNOSIS: Males recognized among other BPE as long as basal plate (pl. 22E); area of Tanindrazanus species by the dark brown to endosomal struts–DPS fusion ovate (pl. 21H). black coloration with pale yellow markings, ETYMOLOGY: The species epithet is an adjecsynthlipsis about 1.5 times the width of an eye, tive in the nominative case and is named for the and eyes not reaching the dorsal and ventral blackish coloration of the body. head margins. This species is similar to T. tene- DISTRIBUTION: Antsiranana and Toamasina bricus, but the larger width of the synthlipsis, provinces (map 8). eyes not reaching the dorsal and ventral head DISCUSSION: Coloration is variable with the margins, and intersegmental sutures carinulate clypeus, postclypeus, coxae, intersternal sutures, between sternites II–IV distinguishes T. margin- median longitudinal depression on sternites, and atus from T. tenebricus . medial sternites pale yellow in some specimens. DESCRIPTION: Medium body size (length: HOLOTYPE: Male: Madagascar: Antsiranana: 12.63 mm, holotype; 11.78–12.63 mm). COL- Sakalava Beach, dwarf littoral forest, 12.26277 ° S ORATION: Dark brown to black with mandibu- 49.3975 ° E, 10 m, 31 May 2001 – 07 Jun 2001, M. lar plates dorsally, postocular dorsally except Irwin, R. Harin’Hala (00006712) (CAS).

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PARATYPES: Madagascar: Antsiranana: Forêt the hind legs. Tanindrazanus notatus is similar to Ambato, 26.6 km 33° NE Ambanja, 13.46444 ° S T. andohahela and T. bemaraha , but the black 48.55166 ° E, 150 m, 08 Dec 2004, B.L. Fisher, 13

apical half of the hind femur and tibia and the

(00044939) (CAS). Montagne Français, 12.325 ° S 49.33333 ° E, 150 m, 15 Feb 2001 – 06 Mar 2001, M. subequal lengths of labial segments II and III dif- Irwin, R. Harin’Hala, 13 (00006705) (SU), 13 ferentiate T. notatus from these species. (00006717) ( USNM) ; 06 Mar 2001 – 20 Mar 2001, M. DESCRIPTION: Medium body size (length: Irwin, R. Harin’Hala, 13 (00045437) ( UCR). Parc 10.17 mm, holotype; 10.17–12.24). COLOR- National Montagne d’Ambre, 12.51444 ° S 49.18138 ° E, ATION: Orange with apical half of scape, pedi- 960 m, 26 Jan 2001 – 29 Jan 2001, M.E. Irwin, E.L. Sch-

cel, antennal segments III–VI, apical half of hind

linger, R. Harin’Hala, 33 (00006715, 00007165, 00007195) (CAS), 13 (00007231) (AMNH); 11 Feb femur, and hind tibia, except basally, dark brown. 2001– 12 Feb 2001, M. Irwin, R. Harin’Hala, 23 Apex of corium, membrane, and distal wing (00006727, 00045465) (CAS); 04 Mar 2001 –19 Mar veins brown. VESTITURE: As in generic descrip- 2001, M. Irwin, R. Harin’Hala, 43 (00006704, tion. STRUCTURE: HEAD: Ventrally with shal- 00006710, 00006726, 00045231) (CAS), 13 (000045328) low anteromedial depression; clypeal apex (BMNH), 13 (000045436) (MNHN), 13 (00045500) (SU). Parc National Montagne d’Ambre, 12.51666 ° S dorsally elevated relative to labrum (pl. 13G, H); 49.18333 ° E, 975 m, 25 Jan 2001 – 11 Feb 2001, M.E. postclypeus with relatively deep, broad medial Irwin, E.L. Schlinger, R. Harin’Hala, 13 (00005368) longitudinal depression to middle of interocular (UCR); 25 Jan 2001 – 29 Jan 2001, M.E. Irwin, E.L. Sch- area; synthlipsis about 1.5 times width of eye; linger, R. Harin’Hala, 23 (00007047, 00007134) (CAS); interocular sulcus near hind margin of eye (pl. 11 Feb 2001 – 04 Mar 2001, M.E. Irwin, E.L. Schlinger,

13B); postocular narrow in dorsal view (pl. 13A);

R. Harin’Hala, 13 (00006145) (BMNH), 13 (00006718) (MNHN). Parc National Montagne d’Ambre, 12.52027 ° S ocelli large, separated by less than diameter of 49.17916 ° E, 1125 m, 19 Mar 2001 – 15 Apr 2001, M. ocellus, located on distinct median tubercle (pl. Irwin, R. Harin’Hala, 23 (00006714, 00006716) (CAS), 13H); eye about one-fourth of head length, not 13 (00006721) (BMNH), 13 (00006722) (AMNH); 21 reaching dorsal and ventral head surfaces; pedi- Apr 2001– 26 Apr 2001, M. Irwin, R. Harin’Hala, 13 cel about one-fourth longer than scape; labial (00006703) (UCR); 14 May 2001 – 30 May 2001, M. Irwin, R. Harin’Hala, 13 (00006711) (USNM); 30 May segment III subequal to II. THORAX: Anterior 2001– 06 Jun 2001, M. Irwin, R. Harin’Hala, 13 pronotal lobe without distinct anterolateral pro- (00006706) (CAS), 13 (00006713) (MNHN). Sakalava jections; posterior pronotal lobe transversely Beach, dwarf littoral forest, 12.26277 ° S 49.3975 ° E, 10 striated; pronotal transverse furrow continuous; m, 07 Apr 2001 – 22 Apr 2001, M. Irwin, R. Harin’Hala, forefemur moderately incrassate; tarsomeres I 13 (00006708) (SU); 13 May 2001 – 16 May 2001, M.

and II combined subequal to III; hemelytron sur-

Irwin, R. Harin’Hala, 13 (00006708) ( USNM) ; 07 Jun 2001 – 25 Jun 2001, M. Irwin, R. Harin’Hala, 13 passing abdominal apex. ABDOMEN: Apex (00006709) ( AMNH) ; 13 Aug 2001 – 20 Aug 2001, M. rounded; dorsal laterotergite II not expanded; Irwin, R. Harin’Hala, 13 (00045396) ( CAS) ; 20 Aug intersegmental sutures carinulate between ster- 2001– 28 Aug 2001, M. Irwin, R. Harin’Hala, 13 nites II – VI and laterally between VI and VII; (00007143) ( UCR) . Toamasina: Botanic Garden near pygophore process subtriangular in lateral view entrance to Andasibe National Park , 18.92633 ° S

(pl. 20H, I); BPE shorter than basal plate; area of

48.40783 ° E, 1025 m, 08 Oct 2001 – 16 Oct 2001, M. Irwin, R. Harin’Hala, 13 (00006725) (CAS). Unknown: endosomal struts–DPS fusion elongate subquad- 23 (00006707, 00045240) (CAS). rate (pl. 21G).

ETYMOLOGY: The species epithet is a past par-

ticiple in the nominative case and is named for

Tanindrazanus notatus , new species the black markings on the hind legs.

Plates 3, 6, 10, 21G; map 6 DISTRIBUTION: Beroboka village from the

DIAGNOSIS: Males are recognized among Toliara province (map 6). other species in this genus by the distinct dull DISCUSSION: The two specimens examined orange-brown coloration with black markings on differ in size (10.17 and 12.24 mm) but are mor-

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 87

phologically similar and were collected from the transversely striated; pronotal transverse furrow same locality. continuous; legs slender; tarsomeres I and II

HOLOTYPE: Male: Madagascar: Toliara: combined subequal to III on forelegs, longer on Beroboka village , 45 km NE Morondava, 19.9775 ° S mid and hind legs; hemelytra surpassing abdom- 44.82483 ° E, 131 m, 05 Feb 2009 – 13 Feb 2009, M. inal apex. ABDOMEN: Apex rounded; dorsal lat- Irwin, R. Harin’Hala (00045368) ( CAS). erotergite II not expanded; intersegmental

PARATYPE: Madagascar: Toliara: Beroboka village , sutures carinulate between sternite II – IV and 45 km NE Morondava, 19.9775 ° S 44.82483 ° E, 131 m, laterally between IV – VI ; pygophore process sub- 19 Dec 2008 – 27 Dec 2008, M. Irwin, R. Harin’Hala, 13

triangular in lateral view (pl. 20H, I); BPE longer (00044868) ( CAS).

than basal plate; area of endosomal struts–DPS

fusion ovate.

Tanindrazanus simulans , new species

ETYMOLOGY: The species epithet is an adjec-

Plates 3, 6, 10

tive in the nominative case and is named after

DIAGNOSIS: Males are recognized among

the similarity to T. tenebricus . other Tanindrazanus species by the nearly uni-

DISTRIBUTION: The two known specimens form dark brown to blackish coloration, syn-

were labeled as originating from “Ivondro.” There thlipsis about one and one-fourth times wider

are at least four localities in the Fianarantsoa, than the width of an eye, eyes reaching ventral

Toamasina, and Toliara provinces that include and almost reaching the dorsal head margins,

the name “Ivondro”; we were unable to deter- and a continuous pronotal transverse furrow.

mine the exact locality. This species is similar to T. tenebricus and T.

HOLOTYPE: Male : Madagascar: Unknown: nigripes , but is distinguished by the width of

Reg. S.E. Ivondro, 1939, A. Seyrig ( MNHN). synthlipsis, eyes reaching the ventral head mar- PARATYPE: Madagascar: Unknown: Reg. S.E. Ivongin but not the dorsal margin, and the pedicel dro, Feb 1940, A. Seyrig, 13 ( MNHN). about one-fifth longer than scape.

DESCRIPTION: Medium body size (length: Tanindrazanus tenebricus , new species 15.40 mm, holotype; 15.10–15.40 mm). COLOR- Plates 3, 6, 10, 18B; map 8

ATION: Blackish except postocular part of the DIAGNOSIS: Males are recognized among head around ocellar tubercle and tarsi pale other Tanindrazanus species by the blackish colbrown. VESTITURE: As in generic description. oration with pale markings on the head and pro-

STRUCTURE: HEAD: Ventrally with shallow notum, synthlipsis about the width of an eye, and anteromedial depression; clypeal apex dorsally large eyes reaching only dorsal head. This species elevated relative to labrum (pl. 13G, H); postclyp- is similar to T. nigripes and T. simulans , from eus with relatively deep, broad medial longitudi- which it differs by the smaller synthlipsis width nal depression to middle of interocular area (pl. and eyes reaching the dorsal but not the ventral 13B); synthlipsis about one and one-fourth times head surface. Tanindrazanus tenebricus is further width of eye; interocular sulcus near hind mar- differentiated from T. nigripes by the interseggin of eye (pl. 13B); postocular narrow in dorsal mental sutures carinulate between sternite II–IV view (pl. 13A); ocelli large, separated by less than and laterally between IV–VI. diameter of ocellus, located on distinct median DESCRIPTION: Medium body size (length: tubercle (pl. 13H); eye about one-third of head 16.37 mm, holotype; 15.40–17.02 mm). COLlength, reaching ventral but not dorsal head sur- ORATION: Blackish with mandibular plates face; pedicel about one-fifth longer than scape; dorsally, postocular dorsally except ocellar labial segment III subequal to II. THORAX: Ante- tubercle, posterolateral margin of pronotum, rior pronotal lobe with very small anterolateral and tarsi pale yellow. Labium light brown, projections (pl. 15B); posterior pronotal lobe except segment II basally, and antennal segment

88 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

VI light brown and segment VII white. VESTI- PARATYPES: Madagascar: Fianarantsoa: 50 km S of TURE: As in generic description. STRUC- Farafangana, Mahabo Mananivo, Ampitavananima For- TURE: HEAD: Ventrally with shallow est, 23.12983 ° S 47.717 ° E, 34 m, 10 Feb 2007 –17 Feb

2007, M. Irwin, F. Parker, R. Harin’Hala, 13 (00006723) anteromedial depression; clypeal apex dorsally (CAS); 03 Mar 2007 – 10 Mar 2007, M. Irwin, F. Parker, elevated relative to labrum (pl. 13G, H); R. Harin’Hala, 13 (00006720) (SU); 17 Mar 2007 –24 postclypeus with relatively deep, broad medial Mar 2007, M. Irwin, F. Parker, R. Harin’Hala, 13 longitudinal depression to middle of interocular (00006724) (UCR). Manombo Special Reserve camp area; synthlipsis about width of eye; interocular site, 32 km SSE of Farafangana, 23.02183 ° S 47.72 ° E, 36

m, 10 Oct 2004 – 21 Oct 2004, M. Irwin, R. Harin’Hala, sulcus near hind margin of eye (pl. 13H); post-

23 (00007141, 00044862) (CAS); 23 Nov 2004 –05 Dec ocular narrow in dorsal view (pl. 13A); ocelli 2004, M. Irwin, R. Harin’Hala, 13 (00044995) large, separated by less than diameter of ocellus, (AMNH); 05 Jan 2005 – 16 Jan 2005, M. Irwin, R. located on distinct median tubercle (pl. 13H); Harin’Hala, 23 (00007046, 00045656) (CAS); 16 Jan eye about one-third of head length, reaching 2005– 20 Jan 2005, M. Irwin, R. Harin’Hala, 23 dorsal but not ventral head surface; pedicel (00045488, 00045723) (BMNH); 13 Mar 2005 –26 Mar

2005, M. Irwin, R. Harin’Hala, 13 (00045301) (UCR); about one-fourth longer than scape; labial seg- 10 Apr 2005 – 24 Apr 2005, M. Irwin, R. Harin’Hala, 23 ment III subequal to II. THORAX: Anterior pro- (00044873, 00045022) (CAS), 13 (00045267) (AMNH); notal lobe with very small anterolateral 22 May 2005 – 06 Jun 2005, M. Irwin, R. Harin’Hala, 73 projections (pl. 15B); posterior pronotal lobe (00007118, 00045270, 00045399, 00045403, 00045496, transversely striated; pronotal transverse furrow 00045606, 00045721) (CAS); 10 Aug 2005 –25 Aug

2005, M. Irwin, R. Harin’Hala, 23 (00044828, divided by paramedian longitudinal ridges (pl.

00045684) (MNHN); 11 Sep 2005 – 25 Sep 2005, M. 15B); legs slender; tarsomeres I and II com- Irwin, R. Harin’Hala, 13 (00045104) (CAS), 13 bined subequal to III on forelegs, longer on mid (00045196) (USNM); 25 Sep 2005 – 02 Oct 2005, M. and hind legs; wings spread out in specimens. Irwin, R. Harin’Hala, 13 (00007101) (SU), 13 ABDOMEN (pl. 18B): Apex rounded; dorsal lat- (00045477) (UCR). Unknown: Reg. S.E. Ivondro, Dec erotergite II not expanded; intersegmental 1938, A. Seyrig, 13 (MNHN); 1939, A. Seyrig, 13

(MNHN). sutures carinulate between sternite II–IV and laterally between IV–VI; pygophore process subtriangular in lateral view (pl. 20H, I); BPE Tanindrazanus varicolor , new species longer than basal plate; area of endosomal Plates 3, 6, 7, 10, 13H, 16A, 17C, 20I, 21H; struts–DPS fusion ovate (pl. 21H). map 7 ETYMOLOGY: The species epithet is an adjec- DIAGNOSIS: Males are recognized among tive in the nominative case and is named for the other species in this genus by the distinct black dark coloration of the body. and orange coloration, labial segments II and III DISTRIBUTION: Specimens are known from subequal in length, and the anterior pronotal the southeastern region of the Fianarantsoa lobe with very small anterolateral protuberances. province (map 8). Two specimens were labeled This species is similar to T. irwini , but differs by as originating from “Ivondro.” There are at least the smaller body size, anteromedial depression four localities in the Fianarantsoa, Toamasina, on the ventral head surface, and intersegmental and Toliara provinces that include the name sutures carinulate between sternite II–VI and lat- “Ivondro”; we were unable to determine the erally between VI and VII, as well as the diagexact locality. nostic features mentioned. HOLOTYPE: Male: Madagascar: Fianarant- DESCRIPTION: Medium body size (length: 15.97 soa: Manombo Special Reserve camp site, 32 km mm, holotype; 15.89–16.12 mm). COLORATION: SSE of Farafangana, 23.02183 ° S 47.72 ° E, 36 m, 16 Dark brown-black with bright orange head, scape, Jan 2005– 20 Jan 2005, M. Irwin, R. Harin’Hala labium, pronotum, scutellum medially, stridula- (00045483) (CAS). tory groove, meso- and metasternal longitudinal

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 89

depressions, hemelytron except posteromedial PARATYPES: Madagascar: Antsiranana: 7 km N of spot, femora except medially on mid leg, tibiae, Joffreville, 12.33333 ° S 49.25 ° E, 360 m, 20 Mar 2001 –07 tarsi, anterior half of laterotergites, sternite III Apr 2001, M. Irwin, R. Harin’Hala, 13 (00006287)

(CAS); 27 Apr 2001 – 13 May 2001, M. Irwin, R. medially, and sternites II – VII anterolaterally. Distal Harin’Hala , 13 (00006288) ( UCR) ; 13 May 2001 –16 half of antennal segment VII white. VESTITURE: May 2001, M. Irwin, R. Harin’Hala, 13 (00006484) As in generic description. STRUCTURE: HEAD ( USNM). Montagne Français, 12.325 ° S 49.33333 ° E, 150 (pl. 13H): Ventrally with shallow anteromedial m, 30 Jan 2001 – 15 Feb 2001, M. Irwin, R. Harin’Hala , depression ; clypeal apex dorsally elevated relative 13 (00006279) (SU); 15 Feb 2001 – 06 Mar 2001, M.

Irwin, R. Harin’Hala, 23 (00006274, 00006281) (CAS); to labrum; postclypeus with relatively deep, broad

06 Mar 2001 – 20 Mar 2001, M. Irwin, R. Harin’Hala, medial longitudinal depression to middle of 13 (00006259) (UCR). Parc National Montagne interocular area; synthlipsis about 1.5 times width d’Ambre, 12.52027 ° S 49.17916 ° E, 1125 m, 11 Feb 2001 – of eye; interocular sulcus near hind margin of eye 04 Mar 2001, M. Irwin, R. Harin’Hala, 23 (00006254, (pl. 13B); postocular narrow in dorsal view (pl. 00006264) (CAS); 19 Mar 2001 – 15 Apr 2001, M. Irwin, 13A); ocelli large, separated by less than diameter R. Harin’Hala, 33 (00006247, 00006256, 00006257)

(CAS), 13 (00006282) (MNHN); 05 Apr 2001 –21 Apr of ocellus, located on distinct median tubercle; eye 2001, M. Irwin, R. Harin’Hala, 13 (00006136) about one-fourth of head length, not reaching dor- (BMNH), 13 (00006267) (AMNH); 21 Apr 2001 –26 sal and ventral head surfaces; pedicel about one- Apr 2001, M. Irwin, R. Harin’Hala, 23 (00006482, fourth longer than scape; labial segment III 00006483) (UCR). Parc National Montagne d’Ambre, subequal to II. THORAX: Anterior pronotal lobe 12.61666 ° S 49.15 ° E, 1116 m, Dec 1968, J. Vadon and A.

Peyrieras, 13 (MNHN). Parc National Montagne with very small anterolateral projections (pl. 15B);

d’Ambre, 12.51666 ° S 49.18333 ° E, 975 m, 25 Jan 2001 – posterior pronotal lobe transversely striated; pro- 11 Feb 2001, M.E. Irwin, E.L. Schlinger, R. Harin’Hala, notal transverse furrow divided by paramedian 23 (00006255, 00006486) (CAS); 04 Mar 2001 –19 Mar longitudinal ridges (pl. 15B); legs slender; tarso- 2001, M.E. Irwin, E.L. Schlinger, R. Harin’Hala, 13 meres I and II combined subequal to III on fore- (00006266) (UCR). Parc National Montagne d’Ambre, and mid legs, longer on hind legs; hemelytron 12.51444 ° S 49.18138 ° E, 960 m, 23 Jan 2001, M.E. Irwin,

23 (00006116, 00006258) (USNM); 26 Jan 2001 –29 Jan reaching or surpassing abdominal apex. ABDO- 2001, M.E. Irwin, E.L. Schlinger, R. Harin’Hala, 23 MEN: Apex rounded; dorsal laterotergite II not (00006253, 00006272) (CAS), 13 (00007065) (SU); 11 expanded; intersegmental sutures carinulate Feb 2001– 12 Feb 2001, M. Irwin, R. Harin’Hala, 23 between sternite II–VI and laterally between VI (00006277, 00006278) (UCR); 04 Mar 2001 –19 Mar and VII; pygophore process subtriangular in lateral 2001, M. Irwin, R. Harin’Hala, 33 (00006252,

00006275, 00006276) (CAS), 13 (00006292) (AMNH); view (pl. 20I); BPE as long as basal plate; area of

19 Mar 2001 – 05 Apr 2001, M. Irwin, R. Harin’Hala, 33 endosomal struts–DPS fusion ovate (pl. 21H). (00006289, 00006291, 00006702) ( CAS). Sakalava ETYMOLOGY: The species epithet is an adjec- Beach, dwarf littoral forest, 12.26277 ° S 49.3975 ° E, 10 tive in the nominative case and is named for the m, 15 Feb 2001 – 06 Mar 2001, M. Irwin, R. Harin’Hala , variable color forms of this species. 13 (00006283) ( CAS) ; 06 Mar 2001 – 20 Mar 2001, M. DISTRIBUTION: Toamasina and the northern Irwin, R. Harin’Hala , 13 (00006260) ( CAS) ; 20 Mar

2001– 07 Apr 2001, M. Irwin, R. Harin’Hala, 13 region of the Antsiranana provinces (map 7).

(00006273) (BMNH); 07 Apr 2001 – 22 Apr 2001, M. DISCUSSION: Some specimens are predomi- Irwin, R. Harin’Hala, 13 (00006265) (MNHN); 13 May nately orange in coloration while others have 2001– 16 May 2001, M. Irwin, R. Harin’Hala, 13 reduced to absent orange markings on the head, (00006498) (AMNH); 31 May 2001 – 07 Jun 2001, M. thorax, scutellum, femora, tibiae, and abdomen. Irwin, R. Harin’Hala, 13 (00006251) (SU); 13 Aug

2001– 20 Aug 2001, M. Irwin, R. Harin’Hala, 13 HOLOTYPE: Male : Madagascar: Antsiranana:

90 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

09 Apr 2001 – 23 Apr 2001, M. Irwin, R. Harin’Hala, 13 sal and ventral head surfaces; pedicel about one- (00006115) (UCR). Botanic Garden near entrance to fourth longer than scape; labial segment III Andasibe National Park, 18.92633 ° S 48.40783 ° E, 1025 subequal to II. THORAX: Anterior pronotal lobe m, 31 Jul 2001 – 15 Aug 2001, M. Irwin, R. Harin’Hala,

with very small anterolateral projections (pl. 15B);

13 (00006250) (UCR); 08 Oct 2001 – 16 Oct 2001, M. Irwin, R. Harin’Hala, 13 (00006246) (BMNH), 13 posterior pronotal lobe smooth; pronotal trans- (00006248) (MNHN), 53 (00006249, 00006280, verse furrow continuous; legs slender; tarsomeres 00006284–00006286) (CAS), 13 (00006290) (SU); 07 I and II combined subequal to III; wings spread Nov 2001– 16 Nov 2001, M. Irwin, R. Harin’Hala, 13 out in specimen. ABDOMEN: Apex rounded; dor- (00005360) (CAS), 13 (00006268) (AMNH).

sal laterotergite II very slightly expanded; interseg-

Unknown: 13 (00006270) (CAS).

mental sutures carinulate between sternites II–V;

pygophore process subtriangular in lateral view

Tanindrazanus vohiparara , new species (pl. 20H, I); BPE shorter than basal plate; endo-

Plates 3, 7, 10; map 8 somal struts ventrally obscured by semiextended

DIAGNOSIS: Males are recognized among endosoma; area of endosomal struts–DPS fusion other species Tanindrazanus by the blackish col- ovate (pl. 21H). oration with pale yellow markings, large eyes ETYMOLOGY: The species epithet is a noun in reaching dorsal and ventral head margins, apposition and is named after the type locality smooth posterior pronotal lobe, a very slightly that is part of Ranomafana National Park, expanded dorsal laterotergite II, and the inter- Madagascar. segmental sutures carinulate between sternites DISTRIBUTION: Ranomafana National Park in II–V. This species is similar to T. nigripes , par- the Fianarantsoa province (map 8). ticularly a variant of this species, but is distin- HOLOTYPE: Male: Madagascar: Fianarantguished by the eye about half of head length, soa: Parc National Ranomafana, Vohiparara, at smooth posterior pronotal lobe, and expanded broken bridge, 21.22616 ° S 47.36983 ° E, 1100 m, dorsal laterotergite II. 04 Mar 2002 – 12 Mar 2002, M. Irwin, R.

DESCRIPTION: Medium body size (length: Harin’Hala (00006139) (CAS). 11.20 mm, holotype). COLORATION: Black with PARATYPE: Madagascar: Fianarantsoa: Parc pale yellow scape basally, antennal segment VII, National Ranomafana, radio tower at forest edge, labial segment III and VI, thoracic sterna except 21.251 ° S 47.40716 ° E, 1130 m, 23 Oct 2005 –30 Oct

2005, M. Irwin, R. Harin’Hala, 13 (00045227) (CAS).

medial longitudinal depression on meso- and metasterna, coxae, trochanters, forefemur ventrally and basally, mid and hind femora except Toliarus , new genus medially, foretibia ventrally, anterior margins of Plates 3, 7, 10, 13I, 19G, 20J, 21I, 22F; map 9 laterotergites, and abdominal sternites except TYPE SPECIES: Toliarus trichrous , new large lateral spots on II–VI and apex of VII. VES- species.

TITURE: As in generic description. STRUC- DIAGNOSIS: Males recognized by a combina-

TURE: HEAD: Ventrally with shallow anteromedial tion of the following characters: the small body depression; clypeal apex slightly elevated relative size, ocelli on a shallow median tubercle, large to labrum (pl. 13G, H); postclypeus with deep, antennal shield that does not conceal antennal broad medial longitudinal depression to middle of insertion in lateral view, 6-segmented antenna, interocular area; synthlipsis about width of eye; and punctate posterior pronotal lobe. This genus interocular sulcus near hind margin of eye (pl. is very similar to Tanindrazanus , from which it 13B); postocular narrow in dorsal view (pl. 13A); differs by the smaller body size, 6-segmented ocelli large, separated by less than diameter of antenna, and punctate posterior pronotal lobe. ocellus, located on distinct median tubercle (pl. DESCRIPTION: MALE: Macropterous, small 13H); eye about half of head length, reaching dor- body size. COLORATION: Red and black or

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 91

orange and black. VESTITURE: Dense, semierect parts of M and Cu veins separate (pl. 17A, C); disto erect, long pale to dark brown setae on head, tal part of M extending beyond apical junction of thorax, legs, corium, and abdomen; setae on tibiae M+Cu (pl. 17A, C); base of M+Cu cell shorter stouter and denser near apex. STRUCTURE: than or as wide as Cu+1A cell (pl. 17B, C). ABDO- HEAD (pl. 13I): Subtriangular in lateral view, lon- MEN: Dorsal laterotergite II not expanded; sterger than wide in dorsal view, shorter than prono- nites shallowly depressed medially (pl. 18C), tum; anteocular region longer than postocular; intersegmental sutures carinulate; spiracles ovoid; clypeal apex not elevated relative to labrum; max- pygophore (pls. 19G, 20J) process subtriangular in illary plate not reaching dorsal surface of clypeus lateral view, directed dorsoposteriad, not surpass- [except nearly reaching in a some specimens of T. ing posterior margin of pygophore; DPS apex trichrous ]; postclypeus depressed (pl. 13B); post- rounded (pl. 21I); endosomal struts reaching posocular broad in dorsal view (pl. 13B); ocelli pres- terior margin of DPS, with anterior and posterior ent, located on shallow median tubercle; distinct ventral processes (pl. 22G); endosoma medially constriction between postocular and neck; gula sclerotized (pl. 21I).

moderately swollen ventrolaterally, not distinctly ETYMOLOGY: This genus is named after the produced beyond ventral head margin; antennal locality province of the type species, Toliara, shield not concealing antennal insertion in lateral Madagascar. The gender is masculine .

view; antenna inserted dorsally on head; scape DISTRIBUTION: Species are known from the surpassing clypeal apex, pedicel slightly curved; southern region of the Toliara province and from flagellum subdivided into BFLA and DFLA; BFLA the Antsiranana province and in habitats between divided into two pseudosegments; DFLA divided 35–180 m elevation. Macrohabitats are described into two pseudosegments; antenna thus appearing as gallery forests, spiny forests, dry deciduous 6-segmented; labium slender; labial segment III forests, and transitional forests.

subequal to II, ventrally straight. THORAX: Prono- DISCUSSION: Female specimens are unknown tum wider than long, anterior margin distinctly for species currently in this genus. The size of the concave, collar distinct (pl. 15B); anterior pronotal maxillary plates is polymorphic in T. trichrous ; lobe shorter than posterior lobe (pl. 15B), more in approximately 40% of the specimens, the maxthan half as wide as posterior lobe, relatively illary plates nearly reach the dorsal clypeal sursmooth; posterior pronotal lobe weakly punctate; face when viewed laterally (pl. 13H).

pronotal longitudinal furrow reaching anterior

but not posterior margin, foveate posteriorly (pl. IDENTIFICATION KEY TO THE MALES OF 15B); pronotal transverse furrow distinct (pl. SPECIES OF Toliarus

15B); lateral depressions on pronotum distinct (pl.

15B), transversely striated; scutellum with two 1. Orange with blackish markings, ocelli sepabroadly separated apical processes, disc medially rated by a distance about the diameter of an depressed; meso- and metasterna shallowly sepa- ocellus, medium body size (> 10 mm).....

rated by transverse suture (pl. 15F); mesosternum.................... karinae , new species with medial and paramedial longitudinal depres- – Tricolored (red and black with dark brown sions; metasternum medially longitudinally forewing membrane), ocelli separated by depressed; MGE with deeply depressed meshlike more than diameter of an ocellus; small cuticle that extends dorsally in lateral view (pl. body size (<10 mm) trichrous , new species 16A); fossula spongiosa on fore- and mid tibiae;

corium restricted to areas adjacent to basal wing Toliarus karinae , new species

veins, with pterostigmalike appearance on antero- Plates 3, 7; map 9

distal margin (pl. 17A, C); distal part of R present DIAGNOSIS: Males are similar to T. trichrous , but not forming cell with M (pl. 17B, C); proximal but is easily distinguished by the larger body size,

92 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

distinct orange body with blackish markings on smaller body size, red and black coloration, and the hemelytra and tibiae, medium sized ocelli, and more punctate posterior pronotal lobe. Males are weak punctuation on the posterior pronotal lobe. most similar to Tanindrazanus anjozorobeus and

DESCRIPTION: Medium body size (length: Tanindrazanus kathrynae , but the smaller body 12.10 mm, holotype). COLORATION: Orange size, red anteocular region and abdominal sterwith antenna except scape ventrally, clavus, fore- nites (except black pygophore), black postocular wing membrane, and apices of tibiae blackish. region, 6-segmented antenna, and punctate pos- Antennifer apically white. VESTITURE: As in terior pronotal lobe distinguishes T. trichrous generic description. STRUCTURE: HEAD: Ven- from these species.

trally with shallow anteromedial depression; DESCRIPTION: Small body size (length: 7.22 postclypeus with relatively deep, narrow medial mm, holotype; 7.22–8.97 mm). COLORATION: longitudinal depression to middle of interocular Red with black head ventrolaterally below eye, area; synthlipsis about two times width of eye; postocular except ventrolateral spot, neck except interocular sulcus near hind margin of eye (pl. dorsomedially, anterior pronotal and propleural 13B); ocelli medium sized, separated by a dis- lobes, prosternum, medial pronotal longitudinal tance about the diameter of an ocellus; eye about furrow on posterior pronotal lobe, meso- and one-fourth of head length, not reaching dorsal metapleura, scutellum, apical half of hind femur, and ventral head surfaces; pedicel about one- hind tibia medially, abdominal sternite VII apifourth longer than scape. THORAX: Anterior pro- cally, and pygophore. Hemelytron dark brown, notal lobe without distinct anterolateral except red corium. VESTITURE: As in generic projections; pronotal transverse furrow divided description. STRUCTURE: HEAD (pl. 13I): Venby paramedian longitudinal ridges (pl. 15B); trally with shallow anteromedial depression; forefemur incrassate; tarsomeres I and II sub- postclypeus with relatively deep, narrow medial equal to III; hemelytron surpassing abdominal longitudinal depression to middle of interocular apex. ABDOMEN: Apex rounded; intersegmental area; synthlipsis about two times width of eye; sutures carinulate between sternites II–VI and interocular sulcus near hind margin of eye (pl. laterally between VI and VII; pygophore process 13B); ocelli small, separated by more than diamsubquadrate in caudal view, with truncate apex; eter of ocellus; eye about one-fourth of head BPE as long as basal plate (pl. 22F); area of endo- length, not reaching dorsal and ventral head sursomal struts–DPS fusion elongate subquadrate faces; pedicel about one-fourth longer than (pl. 21I). scape. THORAX: Anterior pronotal lobe without

ETYMOLOGY: The species epithet is a noun in distinct anterolateral projections; pronotal transthe genitive case and is named after the second verse furrow divided by paramedian longitudinal author’s wife, Karina Wieczorek. ridges (pl. 15B); forefemur incrassate; tarsomeres

DISTRIBUTION: The only known specimen I and II subequal to III; hemelytron surpassing was collected from the Ankarana Reserve in the abdominal apex. ABDOMEN: Apex rounded; Antsiranana province (map 9). intersegmental sutures carinulate between ster-

HOLOTYPE: Male: Madagascar: Antsiranana: nites II – VI and laterally between VI and VII; Ankarana Reserve , 12.91688 ° S 49.14355 ° E, pygophore process subtriangular in lateral view 155m, no date provided, J. Vadon and A. Peyri- (pl. 20J), apex rounded; BPE as long as basal eras ( MNHN). plate (pl. 22F); area of endosomal struts–DPS

fusion elongate subquadrate (pl. 21I).

Toliarus trichrous , new species ETYMOLOGY: The species epithet is an adjec- Plates 3, 7, 10, 13I, 19G, 20J, 21I, 22F; map 9 tive and is named for the body coloration.

DIAGNOSIS: Males are recognized from the DISTRIBUTION: Specimens were collected other species in this genus, T. karinae , by the from the southern region of Toliara (map 9).

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 93

DISCUSSION: Coloration varies with the poste- 00006510) (MNHN); 04 Feb 2004 – 15 Feb 2004, M. rior pronotal lobe and posterior propleural lobe Irwin, F. Parker, R. Harin’Hala, 13 (00006500) (CAS); either uniformly black, black with red lateral mar- 02 Mar 2004 – 11 Mar 2004, M. Irwin, F. Parker, R.

Harin’Hala, 13 (00006509) ( CAS); 05 Apr 2004 –15 Apr gins, or uniformly red. The hind femur may be 2004, M. Irwin, F. Parker, R. Harin’Hala, 13 (00006503) more extensively black, and the black medial spot ( AMNH); 06 Jun 2004 – 20 Jun 2004, M. Irwin, F. Parker, on abdominal sternite VII varies in size. R. Harin’Hala, 13 (00006489) ( CAS); 11 Jul 2004 –25 HOLOTYPE: Male: Madagascar: Toliara: Beza Jul 2004, M. Irwin, F. Parker, R. Harin’Hala , 33 Mahafaly Reserve , Parcelle II near Bellevue , (00006511, 00006512, 00006521) ( CAS); 07 Aug 2004 –

22 Aug 2004, M. Irwin, F. Parker, R. Harin’Hala , 23 23.68983 ° S 44.5755 ° E, 180 m, 21 Nov 2001 –28 (00006487, 00006488) ( USNM) ; 22 Aug 2004 –01 Sep Nov 2001, R. Harin’Hala (00006163) ( CAS). 2004, M. Irwin, F. Parker, R. Harin’Hala , 23 (00006495, PARATYPES: Madagascar: Toliara: Andohahela 00006496) ( SU) ; 01 Sep 2004 – 08 Sep 2004, M. Irwin, F. Natýl Park, Tsimelahy, Parcelle II, 24.93683 ° S Parker, R. Harin’Hala, 13 (00006508) ( UCR). Berenty 46.62666 ° E, 180 m, 09 Dec 2002 – 16 Dec 2002, M. Special Reserve , 8 km NW Amboasary, 25.021 ° S Irwin, F. Parker, R. Harin’Hala, 13 (00006494) ( CAS) ; 46.3055 ° E, 35 m, 10 Jun 2003 – 19 Jun 2003, M. Irwin, F. 16 Dec 2002 – 17 Dec 2002, M. Irwin, F. Parker, R. Parker, R. Harin’Hala, 13 (00006502) ( CAS) ; 08 Nov Harin’Hala, 23 (00006185, 00006186) ( CAS) ; 15 Feb 2003 – 15 Nov 2003, M. Irwin, F. Parker, R. Harin’Hala , 2003– 26 Feb 2003, M. Irwin, F. Parker, R. Harin’Hala, 13 (00005364) ( CAS) ; 21 Dec 2003 – 01 Jan 2004, M. 53 (00006513–00006517) ( CAS) ; 18 Mar 2003 –28 Mar Irwin, F. Parker, R. Harin’Hala, 33 (00006175, 2003, M. Irwin, F. Parker, R. Harin’Hala , 23 (00006184, 00006189, 00006190) ( MNHN); 24 Mar 2004 –04 Apr 00006519) ( USNM) ; 28 Mar 2003 – 08 Apr 2003, M. 2004, M. Irwin, F. Parker, R. Harin’Hala, 13 (00006491) Irwin, F. Parker, R. Harin’Hala , 23 (00006155, ( SU); 23 Jun 2004 – 04 Jul 2004, M. Irwin, F. Parker, R. 00006497) ( BMNH) ; 19 Apr 2003 – 26 Apr 2003, M. Harin’Hala, 13 (00006157) ( BMNH), 13 (00006158) Irwin, F. Parker, R. Harin’Hala , 23 (00006168, ( CAS); 04 Jul 2004 – 18 Jul 2004, M. Irwin, F. Parker, R. 00006169) ( MNHN) ; 29 Jun 2003 – 10 Jul 2003, M. Harin’Hala, 13 (00006170) ( UCR). Beza Mahafaly Irwin, F. Parker, R. Harin’Hala, 23 (00006183, Reserve, Parcelle II near Bellevue , 23.68983 ° S 00006518) ( AMNH) ; 17 Aug 2003 – 24 Aug 2003, M. 44.5755 ° E, 180 m, 10 Nov 2001 – 21 Nov 2001, M. Irwin, Irwin, F. Parker, R. Harin’Hala, 53 (00006176– R. Harin’Hala , 13 (00006208) ( CAS); 04 Dec 2001 –11 00006180) ( CAS) ; 11 Oct 2003 – 19 Oct 2003, M. Irwin, Dec 2001, M. Irwin, R. Harin’Hala, 13 (00006204) F. Parker, R. Harin’Hala, 13 (00006162) ( SU) ; 15 Jan ( UCR). Beza Mahafaly Reserve, Parcelle I near research 2004– 28 Jan 2004, M. Irwin, F. Parker, R. Harin’Hala, station, 23.6865 ° S 44.591 ° E, 165 m, 21 Nov 2001 –28 13 (00006493) ( UCR). Berenty Special Reserve, 8 km Nov 2001, R. Harin’Hala, 33 (00006154, 00006161, NW Amboasary , 25.00666 ° S 46.30333 ° E, 85 m, 25 Oct 00006188) ( CAS) ; 04 Dec 2001 – 11 Dec 2001, R. 2002– 26 Oct 2002, M. Irwin, F. Parker, R. Harin’Hala , Harin’Hala, 13 (00006213) ( CAS) ; 18 Dec 2001 –25 23 (00006153, 00006156) ( CAS) ; 09 Nov 2002 –16 Nov Dec 2001, R. Harin’Hala, 23 (00006193, 00006203) 2002, M. Irwin, F. Parker, R. Harin’Hala, 13 (00006167) ( SU) ; 22 Feb 2002 – 01 Mar 2002, R. Harin’Hala, 13 ( UCR) ; 30 Nov 2002 – 07 Dec 2002, M. Irwin, F. Parker, (00045647) ( CAS) ; 29 Apr 2002 – 19 May 2002, R. R. Harin’Hala, 13 (00006171) ( CAS) ; 14 Dec 2002 –16 Harin’Hala, 13 (00006205) ( BMNH) ; 28 Jul 2002 – 09 Dec 2002, M. Irwin, F. Parker, R. Harin’Hala , 33 Aug 2002, R. Harin’Hala, 13 (00006160) ( CAS) ; 09 Sep (00006181, 00006182, 00006501) ( AMNH) ; 26 Jan 2002 – 20 Sep 2002, R. Harin’Hala , 13 (00006209) 2003– 05 Feb 2003, M. Irwin, F. Parker, R. Harin’Hala, ( CAS) ; 28 Oct 2002 – 10 Nov 2002, M.E. Irwin, F.D. 13 (00006507) ( SU) ; 24 Apr 2003 – 03 May 2003, M. Parker, R. Harin’Hala, 13 (00006191) ( CAS) ; 10 Nov Irwin, F. Parker, R. Harin’Hala, 13 (00006506) ( CAS) ; 2002– 22 Nov 2002, R. Harin’Hala , 23 (00006199, 25 May 2003 – 04 Jun 2003, M. Irwin, F. Parker, R. 00006215) ( USNM) ; 02 Dec 2002 – 12 Dec 2002, R. Harin’Hala, 13 (00006202) ( USNM) ; 22 Nov 2003 –30 Harin’Hala, 33 (00006150, 00006151, 00006212) Nov 2003, M. Irwin, F. Parker, R. Harin’Hala, 13 ( AMNH) ; 12 Dec 2002 – 17 Dec 2002, R. Harin’Hala, (00006490) ( BMNH) ; 30 Nov 2003 – 07 Dec 2003, M. 13 (00006187) ( CAS) ; 17 Dec 2002 – 20 Dec 2002, R. Irwin, F. Parker, R. Harin’Hala , 13 (00006520) Harin’Hala, 13 (00006214) ( CAS) ; 20 Dec 2002 –24 ( BMNH) ; 07 Dec 2003 – 14 Dec 2003, M. Irwin, F. Dec 2002, R. Harin’Hala, 13 (00006164) ( UCR) ; 24 Parker, R. Harin’Hala, 13 (00006492) ( UCR) ; 21 Dec Dec 2002– 02 Jan 2003, R. Harin’Hala , 23 (00006210, 2003– 02 Jan 2004, M. Irwin, F. Parker, R. Harin’Hala, 00006211) ( CAS) ; 09 Jan 2003 – 23 Jan 2003, R. 23 (00006504, 00006522) ( CAS) ; 13 Jan 2004 –23 Jan Harin’Hala, 13 (00006192) ( CAS) ; 04 Feb 2003 – 16 Feb 2004, M. Irwin, F. Parker, R. Harin’Hala, 23 (00006505, 2003, R. Harin’Hala, 13 (00006194) ( BMNH); 02 Mar

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2003– 13 Mar 2003, R. Harin’Hala, 23 (00006159, abdominal tergites I–III declinate toward poste- 00006165) (CAS); 13 Mar 2003 – 23 Mar 2003, R. rior with remaining tergites oriented horizon- Harin’Hala, 23 (00006197, 00006198) (BMNH); 23 tally. This genus is most similar to Rochonia Mar 2003 – 02 Apr 2003, R. Harin’Hala, 23 (00006086, Distant, 1913, from which it differs by the rela-

00006152) (CAS);

02 Apr 2003 – 10 Apr 2003, R.

Harin’Hala, 33 (00006166) (SU), 33 (00006172– tively smooth head and pronotum, hemelytron 00006174) (CAS); 10 Apr 2003 – 21 Apr 2003, R. reaching or surpassing abdominal apex, and dor- Harin’Hala, 23 (00006206, 00006207) (CAS); 21 Apr sal laterotergite II expanded. 2003– 29 Apr 2003, R. Harin’Hala, 13 (00006196) REDESCRIPTION: MALE: Macropterous, (UCR). Cap Ste Marie Special Reserve, 74 km S of Tsi- medium body size. COLORATION: Shiny

hombe, 25.58766 ° S 45.163 ° E, 37 m, 18 May 2003 – 26 May 2003, M. Irwin, F. Parker, R. Harin’Hala, 13 dark brown to black with pale yellow mark- (00006200) (CAS), 13 (00006201) (UCR). Parc ings. VESTITURE: Sparse to dense, semierect National d’Andohahela, Ihazofotsy Parcelle III, to erect, long pale to dark brown setae on 24.83083 ° S 46.53616 ° E, 80 m, 15 Jan 2003 – 26 Jan 2003, head, thorax, legs, corium, and abdomen; setae M. Irwin, F. Parker, R. Harin’Hala, 13 (00007162) on tibiae stouter and denser near apex. (BMNH), 33 (00045213, 00045297, 00045312)

STRUCTURE: HEAD (pl. 13B, J, K): Ovoid or

(MNHN); 24 Aug 2003 – 03 Sep 2003, M. Irwin, F. Parker, R. Harin’Hala, 13 (00045633) (CAS); 12 Nov subquadrate; shorter than pronotum; clypeal 2003– 23 Nov 2003, M. Irwin, F. Parker, R. Harin’Hala, apex not elevated relative to labrum; maxillary 13 (00045242) (UCR); 17 Dec 2003 – 19 Dec 2003, M. plate not reaching dorsal surface of clypeus; Irwin, F. Parker, R. Harin’Hala, 33 (00007208, postclypeus depressed; distinct constriction 00044928, 00045630) (USNM); 14 Jan 2004 –28 Jan between postocular and neck; gula moderately

2004, M. Irwin, F. Parker, R. Harin’Hala, 13 (00007185) (AMNH), 63 (00044953, 00045131, 00045203, swollen ventrolaterally, not distinctly produced 00045298, 00045597, 00045634) (CAS), 23 (00045700, beyond ventral head margin; ocelli present; 00045726) (SU). Unknown: 13 (00006195) (CAS). antennal shield not concealing antennal insertion in lateral view; scape surpassing clypeal

Toxopus Bergroth, 1905 apex; flagellum subdivided into BFLA and

Plates 3, 4, 7, 10, 11, 12, 13B, 13J, 13K, 14, 15B, DFLA; BFLA divided into two pseudoseg- 19H, 19I, 20K, 20L, 21J–L, 22G, 22H, 23E; ments; DFLA divided into three pseudosegmaps 10–13 ments; antenna thus appearing 7-segmented;

Toxopus Bergroth, 1905 , 49: 374. labial segment III subequal to II. THORAX (pl. 15B): Pronotum wider than long, anterior

TYPE SPECIES: Toxopus politus Bergroth, 1905 . margin distinctly concave, collar distinct; ante-

REVISED DIAGNOSIS: Male recognized by the rior pronotal lobe shorter than posterior lobe, shiny dark brown to blackish coloration with more than half as wide as posterior lobe, relapale yellow markings, meso- and metasterna not tively smooth; pronotal longitudinal furrow completely separated by distinct transverse reaching anterior but not posterior margin of suture, dorsal laterotergite II expanded laterally, pronotum, foveate posteriorly; pronotal trans- and laterotergites transversely bicolored. Females verse furrow distinct; lateral depressions on recognized by the apterous condition, shiny dark posterior pronotal lobe distinct; scutellum brown to blackish coloration sometimes with with two moderately separated apical propale yellow markings, head about as wide as cesses, disc medially depressed; meso- and anterior margin of pronotum, 7-segmented metasterna not completely separated by disantenna with short vestiture, scape as long as dis- tinct transverse suture (pl. 15E); mesosternum tance between anterior margin of eye and apex with medial and paramedial longitudinal of head, eye about half head height, anterior pro- depressions; metasternum medially longitudinotal lobe much longer than posterior lobe, fore- nally depressed; MGE with deeply depressed and mid trochanters with small papillae, and meshlike cuticle that extends dorsally in lateral

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 95

view (pl. 16A); forefemur slightly to strongly ented horizontally, intersegments not incrassate; fossula spongiosa on fore- and mid carinulate; sternites flat; external genitalia tibiae; tarsomeres I and II combined subequal short, platelike (pl. 23E).

to III; corium restricted to areas adjacent to DISTRIBUTION: Species are known from all basal wing veins, with pterostigmalike appear- provinces and in habitats between 7–1360 m ance on anterodistal margin (pl. 17A, C); distal elevation. Macrohabitats include tropical dry forpart of R present but not forming cell with M ests, dwarf littoral forests, low-altitude dense (pl. 17B, C); proximal parts of M and Cu sepa- humid forests, low-altitude littoral rainforests, rate (pl. 17A, C); distal part of M extending high-altitude rainforests, spiny forests, gallery beyond apical junction of M+Cu (pl. 17A, C); forests, sclerophyl forests, montane rainforest, base of M+Cu cell shorter than or as wide as dry deciduous forests, and transitional forests.

Cu+1A cell (pl. 17B, C). ABDOMEN: Dorsal lat- DISCUSSION: Bergroth (1905) described this erotergite II expanded; sternites shallowly genus from a single female specimen. He emphadepressed medially (pl. 18C), intersegmental sized the overall head structure and strongly sutures carinulate; spiracles ovoid; pygophore curved hind tibia as diagnostic for this genus. process subtriangular in lateral view (pl. 20K, However, head shape and structure is variable L), directed dorsoposteriad, apex rounded; among species in some ectrichodiine genera (e.g., DPS apex rounded (pl. 21J–L). FEMALE: Dif- Microstemmatoides Putshkov, 1985 , Daraxa Stål , fers from males in the following characteris- 1859, Gibbosella ) and the leg armature can vary tics: apterous; reduced yellow markings; head among and within species. As examples, the preslarger, about as wide as anterior pronotal mar- ence and location of patches of papillae on the gin, shorter or longer than pronotum; femora and trochanters may differ among species postclypeus not depressed (pl. 13A); antennal of Rhiginia Stål, 1859 , males and females of the shield not expanded; antennal vestiture much same species in Racelda Signoret, 1863 , and aptershorter; scape as long as distance between ous and. macropterous forms within species of anterior eye margin and apex of head; ocelli Glymmatophora . Similarly, femoral and tibial absent; eyes small, about half head height; spines may vary in presence, size, or number in labial segments II shorter than or subequal to species of Glymmatophora . Finally, the curvature segment III; collar barely distinct; anterior of the hind tibia can vary between species in a pronotal lobe longer than and as wide as pos- genus (straight or slightly curved in Pothea Amyot terior lobe (pl. 15A), without distinct antero- and Serville. 1843, Katanga Schouteden, 1903 , lateral projections; pronotal longitudinal Centraspis Schaum, 1862 ) and within species furrow reduced to deep medial depression (straight or slightly curved in females of Ectrichonear posterior margin of anterior pronotal dia crux [Thunberg] 1783). Curvature and robustlobe (pl. 15A); lateral depressions on posterior ness of the hind tibia in females of Toxopus politus pronotal lobe obsolete; scutellum weakly Bergroth, 1905, appears to be fairly extreme, and developed; meso- and metasterna platelike we found this character variable in two female with shallow, wide medial longitudinal depres- specimens that we assign to Toxopus : one specision; MGE reduced, shallow, not extending men has a more robust and strongly curved hind dorsally in lateral view (pl. 16B, C); fore- and tibia than the other. We are unable to definitively mid trochanters with patches of small papillae assign either specimen to T. politus based on the (pl. 16D, E); hind femur slightly curved in dor- original description; the holotype of T. politus is sal view; fossula spongiosa larger; dorsal lat- missing and could not be examined for confirmaerotergite II not expanded; abdominal tergites tion (see discussion for T. politus ). Comparison of I– III declinate toward posterior, intersegmen- these two specimens with a female with cylindrical tal sutures carinulate, remaining tergites ori- head and slender, slightly curved hind tibia

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revealed many morphological similarities: apter- IDENTIFICATION KEY TO THE MALES OF ous, shiny blackish coloration, head about as wide SPECIES OF Toxopus

as anterior margin of pronotum, 7-segmented

antenna with short vestiture, scape as long as dis- 1. Head cylindrical (pl. 13J)................ 2 tance between anterior eye margin and apex of – Head subquadrate (pl. 13A) or ovoid (pl. 13F) 9 head, eyes small, and abdominal tergites I– III 2. Head ventrally flat....................... 3 declinate toward posterior, among several other – Head with medial depression either restricted features. Given the abundance of morphological to anteocular region or extending to postsimilarities between these females and known vari- ocular................................ 4 ability in head shape and leg structure in other 3. Pedicel not distinctly curved; sternal interseg- Ectrichodiinae genera, the genus is here rede- mental sutures carinulate between II – IV and scribed to accommodate females with various laterally between IV – VI.. insignis , new species head shapes and slender, less curved, and unarmed – Pedicel slightly curved; sternal intersegmental hind tibia. Based on morphology, geographic sutures carinulate between II and III and latproximity, and molecules, we also associated the erally between III – VI.... fisheri , new species female with the cylindrical head and slender hind 4. Head with ventral anteromedial depression; legs with males of a new species, T. griswoldi , postocular broad in dorsal view (pl. 13B); enabling us to describe Toxopus males for the first sublateral antennal insertion; pronotal transtime. Attempts to associate the other two females verse suture incomplete, divided by paramewith conspecific males based on morphology, dian ridges (pl. 15B)................... 5 geography, and molecules failed. These two females – Head with ventral medial depression extending remain undescribed for the time being since, based to postocular; postocular narrow in dorsal on head shape, presence of a ventral medial depres- view (pl. 13A); dorsal antennal insertion; sion on the head, and geographic proximity, they pronotal transverse suture complete, not could represent females of any of the following divided by paramedian ridges............ species: T. ampitavananima , T. basalis , T. brucei , T. .............. melobrunneus , new species namoroka , T. parkeri , T. signoretii , T. simulans , T. 5. Posterior pronotal lobe smooth; sternal intersteineri, T. tibialis , and T. vazimba . segmental sutures carinulate between II – IV The monotypic genus Rochonia was described and laterally between IV – VI............. by Distant from one macropterous male speci-..................... toliara , new species men from the Seychelles in 1913. The holotype – Posterior pronotal lobe striated; sternal interof R. galeatus Distant, 1913 , which is fairly dam- segmental suture carinulation pattern not as aged and has the ventral surface not visible, was previously mentioned.................. 6 examined at the BMNH. It is clear that Rochonia 6. Synthlipsis width 1.5 times width of eye; stershares many morphological features with Toxo- nal intersegmental sutures carinulate pus, while few characters distinguish the two between II and III and laterally between III genera (see differential diagnosis for Toxopus ). and IV.......... antsiranana , new species Based on these morphological similarities, Toxo- – Synthlipsis width two times width of eye; sterpus may in fact belong to Rochonia . We refrain nal intersegmental suture carinulation patfrom synonymizing Toxopus with Rochonia until tern not as previously mentioned........ 7 additional specimens of Rochonia become avail- 7. Postclypeal depression extending to interocuable that will allow examination of ventral char- lar sulcus; sternal intersegmental sutures acters, antennal segmentation, and intraspecific carinulate between II and III and laterally variation. Future phylogenetic analyses including between III – V. ambohitantely , new species Rochonia and Toxopus may also provide support – Postclypeal depression extending to middle of for synonymy. interocular area (pl. 13B); sternal interseg-

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mental suture carinulation pattern not as nearly reaching ventral head surface in latpreviously mentioned.................. 8 eral view; sternal intersegmental sutures

8. Postclypeal depression relatively broad; ventral carinulate between II – IV and laterally margin of labial segment III convex (pl. 13J, between IV – VI........ basalis , new species D, E, F); sternal intersegmental sutures cari- 15. Posterior pronotal lobe weakly transversely strinulate between II and III and laterally ated; sternal intersegmental sutures carinulate between III – VII.... italaviana , new species between II – VI.......... parkeri , new species

– Postclypeal depression narrow; ventral margin – Posterior pronotal lobe smooth; sternal interof labial segment III straight (pl. 13G–I, K); segmental sutures carinulate between II – IV sternal intersegmental sutures carinulate and laterally between IV – VI............. between II – IV and laterally between IV –..................... brucei , new species VII................ griswoldi , new species 16. Only sternal intersegmental suture between

9. Anteocular region as long as postocular (pl. II – III carinulate...................... 17 13J)................................. 10 – Multiple sternal intersegmental sutures carinu-

– Anteocular region longer than postocular (pl. late................................. 18 13K)................................ 11 17. Synthlipsis about two times width of eye;

10. Postclypeal depression extending to interocu- postocular broad in dorsal view (pl. 13B); lar sulcus; ventral margin of labial segment posterior pronotal lobe smooth.......... III straight (pl. 13G–I, K); posterior pronotal............... miandritsara , new species lobe striated......... pallidus , new species – Synthlipsis about 1.5 times width of eye; post-

– Postclypeal depression extending to middle of ocular narrow in dorsal view (pl. 13A); posinterocular area; ventral margin of labial terior pronotal lobe weakly transversely segment III convex (pl. 13C–F, J); posterior striated............. simulans , new species pronotal lobe smooth................... 18. Synthlipsis at least 1.5 times width of eye; eye................ farafangana , new species not distinctly reaching dorsal head surface

11. Head about as long as wide in dorsal view12.................................... 19

– Head longer than wide in dorsal view..... 16 – Synthlipsis about width of eye; eye reaching or

12. Synthlipsis about width of eye; eye reaching nearly reaching dorsal head surface....... dorsal head surface in lateral view; sternal...... signoretii (Reuter) , new combination intersegmental sutures carinulate between II 19. Synthlipsis about two times width of eye; and III and laterally between III – V....... anterior pronotal lobe with distinct, small................. toamasina , new species anterolateral protuberances (pl. 15B); poste-

– Synthlipsis at least 1.5 times width of eye; eye rior pronotal lobe smooth............. 20 not reaching dorsal head surface in lateral – Synthlipsis about 1.5 times width of eye; anteview; carinulation patterns on the sternal rior pronotal lobe without distinct anterolatintersegmental sutures not as previously eral protuberances; posterior pronotal lobe mentioned........................... 13 weakly transversely striated..............

13. Synthlipsis about 1.5 times width of eye.. 14............ ampitavananima , new species

– Synthlipsis about two times width of eye.. 15 20. Ventral surface of labial segment III straight

14. Head ventrally with anteromedial depression; (pl. 13G–I, K); sternal intersegmental sutures eye not distinctly reaching ventral head sur- carinulate between II – VI and laterally face in lateral view; sternal intersegmental between VI and VII................... 21 sutures carinulate between II – VI......... – Ventral surface of labial segment III convex (pl...................... tibialis , new species 13C–F, J); sternal intersegmental sutures

– Head with medial depression extending along carinulate between II – IV and laterally entire ventral head surface; eye reaching or between IV – VI....... steineri , new species

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21. Interocular sulcus near hind margin of eye chanters, femora except dorsoapical half, fore- (pl. 13B); head, including postclypeal de- and mid tibiae medially, hind tibia subbasally pression, anterior pronotal lobe, and legs and medially, anterior half of laterotergites, reddish brown...... namoroka , new species abdominal sternites medially (except anterior, – Interocular sulcus anterior to hind margin of posterior, and lateral margins), sternites II – VI eye; head dark brown with postclypeal anterolaterally, and lateral margin of sternite depression and labium pale; anterior prono- VII. Antennal segments VI and VII, hemelytal lobe generally dark brown; legs dark tron, and tarsi brown. VESTITURE: Moderbrown and pale..... vazimba , new species ately dense; other features as in generic

description. STRUCTURE: HEAD (pl. 13J): IDENTIFICATION KEY TO THE FEMALES OF Cylindrical in lateral view; longer than wide in

SPECIES OF TOXOPUS dorsal view; ventrally with shallow anterome-

dial depression; anteocular region longer than 1. Hind tibia distinctly curved, armed with small

postocular; postclypeus with deep, narrow papillae (pl. 16D, E); shiny dark brown to

medial longitudinal depression to interocular black; basal part of scape and antennal seg-

sulcus; synthlipsis about two times width of eye; ments V – VII entirely, except basal half of V,

interocular sulcus posterior to hind margin of pale; dorsal surface of femora basally, fore-

eye; postocular broad in dorsal view (pl. 13B); tibia ventrally except basally and apically,

ocelli small, located on shallow median tuberrings on mid and hind tibiae, and tarsi pale

cle, separated by a distance less than or equal to to brown (based on original description)..

diameter of ocellus; eye about one-fourth of

........................ politus Bergroth

head length, not reaching dorsal and ventral – Hind tibia not distinctly curved, unarmed;

head surfaces; antenna inserted sublaterally on shiny blackish with extensive pale to brown

head; pedicel about one-fourth longer than markings........... griswoldi , new species

scape, slightly curved; labium slender; labial

segment III ventrally convex. THORAX: Anterior

Toxopus ambohitantely , new species pronotal lobe with relatively small anterolateral

Plates 3, 7, 10, 13J; map 11 projections (pl. 15B); posterior pronotal lobe DIAGNOSIS: Males are recognized among transversely striated; pronotal transverse furrow other Toxopus species by the moderately dense divided by paramedian longitudinal ridges (pl. vestiture, coloration, elongate cylindrical head in 15B); lateral depressions of posterior pronotal lateral view, relatively deep postclypeal depres- lobe transversely striated; hemelytron surpasssion that reaches the interocular sulcus, sublat- ing abdominal apex. ABDOMEN: Apex rounded, eral antennal insertion, labial segment III sinuate, or medially notched; intersegmental ventrally convex, and intersegmental sutures sutures carinulate between sternites II and III carinulate between sternites II and III and later- and laterally between III – V; pygophore process ally between III – V. This species is similar to T. subtriangular in caudal view (pl. 19I), not suritalaviana, T. griswoldi , T. insignis , T. fisheri , T. passing posterior margin of pygophore; BPE as antsiranana , and T. toliara , but is distinguished long as basal plate; endosomal struts not reachfrom them by the intersegmental sutures carinu- ing posterior margin of DPS (pl. 21L), with late between sternites II and III and laterally anterior and posterior ventral processes (pl. between III – V, subtriangular pygophore process, 22G); area of endosomal struts–DPS fusion and endosoma medially sclerotized. ovate; endosoma medially sclerotized (pl. 21L). DESCRIPTION: MALE: Body length: 9.93 mm ETYMOLOGY: The species epithet is a noun in (holotype), 9.83–10.18 mm. COLORATION: apposition and is named after the type locality, Blackish with pale yellow corium basally, tro- Ambohitantely, Madagascar.

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 99

DISTRIBUTION: Antananarivo, Antsiranana,

Mahajanga, and Toamasina provinces (map 11).

DISCUSSION: In some specimens, the base of

the scape, antennal segments VI and VII, scutel-

lar processes, femora, tibiae, and corium are

more pale yellow than dark brown. The labium

may be light brown.

HOLOTYPE: Male : Madagascar: Antanana-

rivo: 46 km NE of Ankazobe: Ambohitantely,

18.198 ° S 47.2815 ° E, 700 m, 15 Oct 2004 –01 Nov

2004, M. Irwin, F. Parker, R. Harin’Hala

(00006782) ( CAS).

PARATYPES: Madagascar: Antananarivo: 46 km NE

of Ankazobe: Ambohitantely, 18.198 ° S 47.2815 ° E, 700

m, 31 Oct 2003 – 07 Nov 2003, M. Irwin, R. Harin’Hala,

33 (00006788–00006790) ( CAS); 05 Mar 2004 –19 Mar

2004, M. Irwin, R. Harin’Hala, 13 (00006791) ( CAS);

19 Mar 2004 – 28 Mar 2004, M. Irwin, R. Harin’Hala,

13 (00006778) (SU); 15 Oct 2004 – 01 Nov 2004, M.

Irwin, F. Parker, R. Harin’Hala, 33 (00006779–

00006781) ( USNM), 23 (00006783, 00006784) ( CAS),

33 (00006785–00006787) ( MNHN); 01 Nov 2004 –14

Nov 2004, M. Irwin, R. Harin’Hala, 13 (00006792)

( UCR); 17 Apr 2005 – 26 Apr 2005, M. Irwin, R.

Harin’Hala, 23 (00044905, 00045071) ( AMNH), 23

(00045364, 00045369) ( BMNH). Antsiranana: Parc

National Montagne d’Ambre, 12.51666 ° S 49.18333 ° E,

975 m, 04 Mar 2001 – 19 Mar 2001, M.E. Irwin, E.L.

Schlinger, R. Harin’Hala, 13 (00006120) ( CAS). Saka-

lava Beach, dwarf littoral forest, 12.26277 ° S 49.3975 ° E,

10 m, 15 Feb 2001 – 06 Mar 2001, M. Irwin, R. MAP 10. Localities of Toxopus antsiranana , T. gris- Harin’Hala, 13 (00006794) ( USNM); 22 Apr 2001 –27 woldi, T. miandritsara , T. toamasina , T. toliara , and Apr 2001, M. Irwin, R. Harin’Hala, 13 (00045583) T. vazimba .

(SU). Mahajanga: Reserve Speciale de Bemarivo, 23.8

km 223° SW Besalampy, 16.925 ° S 44.36833 ° E, 30 m, 19

tuberances, and intersegmental sutures carinulate Nov 2002– 23 Nov 2002, Fisher et al., 13 (00006793)

( CAS). Toamasina: 7 km SE of Andasibe National Park between sternites II and III and laterally between headquarters, 18.96266 ° S 48.45266 ° E, 1050 m, 07 Jan III – VI. This species is similar to T. vazimba , but 2001– 22 Jan 2001, M. Irwin, R. Harin’Hala, 13 the smaller synthlipsis width, interocular sulcus (00007142) ( CAS). Botanic Garden near entrance to located near the hind margin of the eye, eye one- Andasibe National Park, 18.92633 ° S 48.40783 ° E, 1025 half of the head length in lateral view, anterior m, 01 Sep 2001 – 05 Sep 2001, M. Irwin, R. Harin’Hala,

pronotal lobe lacking distinct anterolateral protu- 13 (00006795) ( MNHN); 19 Sep 2001 – 26 Sep 2001, M.

Irwin, R. Harin’Hala, 13 (00045254) ( UCR). berances, striated posterior pronotal lobe, intersegmental sutures carinulate between sternites II Toxopus ampitavananima , new species and III and laterally between III – VI, and BPE as Plates 3, 7, 10, 22G; map 12 long as the basal plate distinguish T. ampitavana-

DIAGNOSIS: Males are recognized among other nima from T. vazimba .

Toxopus species by the subquadrate head shape in DESCRIPTION: MALE: Body length: 10.67 mm lateral view that is longer than wide in dorsal view, (holotype), 10.50–11.91 mm. COLORATION: anterior pronotal lobe without anterolateral pro- Blackish with pale yellow mandibular and maxil-

100 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

gin of sternite VI, and lateral margin of abdomen

except apex of sternite VII. Labium and remain-

der of pronotum brown. VESTITURE: Moder-

ately dense; other features as in generic

description. STRUCTURE: HEAD: Subquadrate

in lateral view (pl. 13K); longer than wide in dor-

sal view; ventrally with deep medial depression;

anteocular region longer than postocular (pl.

13K); postclypeus with deep, broad medial lon-

gitudinal depression to interocular sulcus; syn-

thlipsis about 1.5 times width of eye; interocular

sulcus near hind margin of eye (pl. 13B); post-

ocular narrow in dorsal view (pl. 13A); ocelli

large, located on shallow median tubercle (pl.

13K), separated by less than diameter of ocellus;

eye about half of head length, not reaching dor-

sal and ventral head surfaces; antenna inserted

dorsally on head; pedicel about one-fourth lon-

ger than scape, slightly curved; labium stout;

labial segment III ventrally straight (pl. 13K).

THORAX: Anterior pronotal lobe without antero-

lateral projections; posterior pronotal lobe

weakly to strongly transversely striated; pronotal

transverse furrow divided by paramedian longi-

tudinal ridges (pl. 15B); lateral depressions of

posterior pronotal lobe weakly to strongly trans-

versely striated; hemelytron reaching or surpass- MAP 11. Localities of Toxopus ambohitantely , T. ing abdominal apex. ABDOMEN: Apex rounded; basalis , T. farafangana , T. melobrunneus , T. parkeri , intersegmental sutures carinulate between ster- and T. tibialis . nites II and III and laterally between III–VI;

pygophore process subtriangular in caudal view lary plates dorsally, postclypeal medial longitudi- (pl. 19I), not surpassing posterior margin of nal depression, anterolateral area anterior of pygophore; BPE as long as basal plate; endointerocular sulcus, scape basally, pedicel basally, somal struts reaching posterior margin of DPS antennal segment III basally, antennal segment (pl. 21J, K), with anterior and posterior ventral IV apically, antennal segments VI and VII, ante- processes (pl. 22G); area of endosomal struts– rior pronotal lobe anterolaterally, posterior mar- DPS fusion elongate subquadrate (pl. 21K); gin of posterior pronotal lobe, stridulatory endosoma medially sclerotized (pl. 21L). process, meso- and metasterna, scutellar pro- ETYMOLOGY: The species epithet is a noun in cesses apically, corium basally, coxae, trochan- apposition and is named after the type locality, ters, forefemur basally and ventrally, mid femur Ampitavananima Forest, Madagascar. basally and subapically, hind femur except medi- DISTRIBUTION: Antsiranana, Fianarantsoa, ally, foretibia except basally, mid tibia except sub- Toamasina, and Toliara provinces (map 12). basally, hind tibia apically, tarsi, anterior half of DISCUSSION: The coloration in this species is laterotergites, abdominal sternites II–V except variable: the pale yellow coloration on the head medially and laterally, ventrolateral anterior mar- and scutellum may be reduced, the anterior pro-

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 101

notal lobe may have more pale yellow coloration

throughout, the posterior pronotal lobe may be

dark brown, and the entire ventromedial abdom-

inal surface may be pale yellow or nearly so.

HOLOTYPE: Male : Madagascar: Fianarant-

soa: 50 km S of Farafangana, Mahabo Mananivo,

Ampitavananima Forest, 23.12983 ° S 47.717 ° E, 34

m, 14 Apr 2007 – 21 Apr 2007, M. Irwin, F. Parker,

R. Harin’Hala (00006768) ( CAS).

PARATYPES: Madagascar: Antsiranana: Parc

National Montagne d’Ambre, 12.51444 ° S 49.18138 ° E,

960 m, 04 Mar 2001 – 19 Mar 2001, M. Irwin, R.

Harin’Hala, 13 (00006719) ( CAS). Fianarantsoa: 50

km S of Farafangana, Mahabo Mananivo, Ampitavana-

nima Forest, 23.12983 ° S 47.717 ° E, 34 m, 13 Jan 2007 –

20 Jan 2007, M. Irwin, F. Parker, R. Harin’Hala, 33

(00006771–00006773) ( CAS); 04 Feb 2007 –10 Feb

2007, M. Irwin, F. Parker, R. Harin’Hala, 33 (00006762–

00006764) ( CAS); 10 Feb 2007 – 17 Feb 2007, M. Irwin,

F. Parker, R. Harin’Hala, 13 (00006766) ( UCR); 17 Feb

2007– 24 Feb 2007, M. Irwin, F. Parker, R. Harin’Hala,

13 (00006740) ( CAS); 10 Mar 2007 – 17 Mar 2007, M.

Irwin, F. Parker, R. Harin’Hala, 43 (00006743–

00006746) ( BMNH), 23 (00006755, 00006756) ( UCR);

17 Mar 2007 – 24 Mar 2007, M. Irwin, F. Parker, R.

Harin’Hala, 13 (00006741) (SU); 24 Mar 2007 –31 Mar

2007, M. Irwin, F. Parker, R. Harin’Hala, 13 (00006769)

( CAS); 07 Apr 2007 – 14 Apr 2007, M. Irwin, F. Parker,

R. Harin’Hala, 43 (00006737–00006739, 00006753)

( MNHN), 13 (00006754) ( CAS); 21 Apr 2007 –29 Apr

2007, M. Irwin, F. Parker, R. Harin’Hala, 13 (00006761) MAP 12. Localities of Toxopus ampitavananima , T. ( CAS); 06 May 2007 – 14 May 2007, M. Irwin, F. Parker, brucei , T. fisheri , T. italaviana , T. pallidus , and T. R. Harin’Hala, 13 (00006770) ( USNM); 14 May 2007 – steineri .

21 May 2007, M. Irwin, F. Parker, R. Harin’Hala, 13

102 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

Site, Analalava 7 km SW of Foulpointe, 17.69333 ° S postocular broad in dorsal view (pl. 13B); ocelli 49.46027 ° E, 18 m, 29 Feb 2008 – 07 Mar 2008, M. Irwin, small, located on shallow median tubercle (pl. R. Harin’Hala, 13 (00044815) ( CAS). Toliara: Forêt

13J), separated by a distance less than or equal

Ivohibe 55.0 km N Tolagnaro, 24.56888 ° S 47.20388 ° E,

200 m, 02 Dec 2006 – 04 Dec 2006, B.L. Fisher et al., 13 to diameter of ocellus; eye about one-fourth of (00006733) ( CAS). Unknown: 13 (00006757) ( CAS). head length, not reaching dorsal and ventral head surfaces; antenna inserted sublaterally on

Toxopus antsiranana , new species head; pedicel about one-fourth longer than

Plates 3, 7, 10; map 10 scape, slightly curved; labium slender; labial seg-

DIAGNOSIS: Males are recognized among ment III ventrally straight (pl. 13K). THORAX: other species in this genus by a combination of Anterior pronotal lobe with small anterolateral the following characters: the coloration, elongate projections (pl. 15B); posterior pronotal lobe cylindrical head in lateral view, synthlipsis 1.5 weakly transversely striated; pronotal transverse times width of an eye, antenna inserted sublater- furrow divided by paramedian longitudinal ally on the head, labial segment III ventrally ridges (pl. 15B); lateral depressions of posterior straight, and intersegmental sutures carinulate pronotal lobe weakly transversely striated; hembetween sternites II and III and laterally between elytron surpassing abdominal apex. ABDOMEN: III and IV. Males are similar to males of T. itala- Apex rounded or sinuate; intersegmental sutures viana, T. griswoldi , T. insignis , T. fisheri , and T. carinulate between sternites II and III and latertoliara, but are distinguished by the smaller ally between III and IV; pygophore process synthlipsis width and intersegmental sutures curved in lateral view (pl. 20K), elongate in caucarinulate between sternites II and III and later- dal view (pl. 19H), surpassing posterior margin ally between III and IV. of pygophore; BPE shorter than basal plate;

DESCRIPTION: MALE: Body length: 12.21 endosomal struts reaching posterior margin of mm (holotype), 11.10–12.21 mm. COLOR- DPS (pl. 21J, K), with anterior and posterior

ATION: Blackish with pale yellow mandibular ventral processes (pl. 22G); area of endosomal plates dorsally, postclypeus between antennifers, struts–DPS fusion elongate subquadrate (pl. antennifer, labial segment IV except basally, 21K); endosoma with sclerotized, hairlike, denscape basally, pedicel basally, base of antennal ticles on posterior margin (pl. 21J, K).

segment III, antennal segment VII, metaster- ETYMOLOGY: The species epithet is a noun in num laterally, corium basally, mid and hind apposition and is named after the Antsiranana coxae posteriorly, trochanters, forefemur basally province in Madagascar, where the type speci- and subapically, mid and hind femora except men was collected.

medially, foretibia medially and subbasally, mid DISTRIBUTION: Toamasina and southern tibia subbasally and subapically, hind tibia region of Antsiranana provinces (map 10). basally and subapically, tarsi, anterior half of lat- DISCUSSION: Coloration varies with the dorsal erotergites, and abdominal sternites medially area between antennifers sometimes brown. and anterolaterally. VESTITURE: Dense ; other HOLOTYPE: Male : Madagascar: Antsiranana: features as in generic description. STRUCTURE: Forêt Ambanitaza, 26.1 km 347° Antalaha,

HEAD: Cylindrical in lateral view (pl. 13J); lon- 14.67944 ° S 50.18361 ° E, 240 m, 26 Nov 2004, B.L. ger than wide in dorsal view; ventrally with Fisher (00044804) (CAS).

shallow anteromedial depression; anteocular PARATYPES: Madagascar: Antsiranana: Forêt region longer than postocular (pl. 13K) ; Ambanitaza, 26.1 km 347° Antalaha, 14.67944 ° S 50.18361 ° E, 240 m, 26 Nov 2004, B.L. Fisher, 13

postclypeus with shallow, narrow medial longi-

(00045129) (CAS). Marojejy National Park, 5 km W

tudinal depression to middle of interocular area; Manantenina village, Camp Mantella, 14.43816 ° S synthlipsis about 1.5 times width of eye; interoc- 49.774 ° E, 490 m, 25 Dec 2004 – 30 Dec 2004, M. Irwin, ular sulcus near hind margin of eye (pl. 13B); R. Harin’Hala, 13 (00044835) (CAS); 05 Jan 2005 –10

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 103

Jan 2005, M. Irwin, R. Harin’Hala, 33 (00006128, VESTITURE: Very sparse; other features as in 00006970, 00007099) (UCR); 25 Feb 2005 –04 Mar generic description. STRUCTURE: HEAD: Sub- 2005, M. Irwin, R. Harin’Hala, 13 (00045652) (CAS); quadrate in lateral view (pl. 13K); as long as wide 18 Mar 2005 – 25 Mar 2005, M. Irwin, R. Harin’Hala,

in dorsal view; ventrally with deep medial

13 (00006098) (CAS); 25 Mar 2005 – 04 Apr 2005, M.

Irwin, R. Harin’Hala , 13 (00006100) (SU) ; 18 May depression ; anteocular region longer than post- 2005– 30 May 2005, M. Irwin, R. Harin’Hala, 13 ocular (pl. 13K) ; postclypeus with deep, broad (00007189) (SU). Toamasina: Fampanambo , 15.37550 ° S medial longitudinal depression to interocular 49.62175 ° E, 108 m, Feb 1959, J. Vadon, 23, ( MRAC) ; sulcus; synthlipsis about 1.5 times width of eye; Feb 1961, J. Vadon, 23, ( MRAC). Ivoloina Zoological

interocular sulcus anterior to hind margin of eye;

Park, 12 km NW of Toamasina, 18.05933 ° S 49.35790 ° E,

29 m, 16 Nov 2000 – 20 Nov 2000, Dolin and Andre- postocular narrow in dorsal view (pl. 13A); ocelli eva, 13 (TLMF). Nosy Mangabe, 15.49324 ° S large, located on shallow median tubercle (pl. 49.76776 ° E, 266 m, 27 Dec 2003, R. Dolin, 13 (TLMF). 13K), separated by less than diameter of ocellus; Périnet (Andasibe), 18.82666 ° S 48.44778 ° E, 1119 m, 19 eye about half of head length, not reaching dor- Dec 2001 – 31 Dec 2001, V. Dolin, 13 (TLMF). sal head surface, reaching ventral head surface; Unknown: 13 (00007191) (CAS).

antenna inserted dorsally on head; pedicel about

one-fourth longer than scape, slightly curved;

Toxopus basalis , new species labium stout; labial segment III ventrally straight

Plates 3, 7, 10, 22H; map 11 (pl. 13K). THORAX: Anterior pronotal lobe with DIAGNOSIS: Males recognized among other very small anterolateral projections (pl. 15B); species in Toxopus by the coloration, sparse ves- posterior pronotal lobe weakly transversely strititure, head as long as wide, synthlipsis 1.5 times ated; pronotal transverse furrow divided by parawidth of an eye, interocular sulcus anterior to the median longitudinal ridges (pl. 15B); lateral hind margin of the eye, eye reaching the ventral depressions of posterior pronotal lobe weakly head margin, striated posterior pronotal lobe, transversely striated; hemelytron surpassing ter- and intersegmental sutures carinulate between gite VII and not reaching, reaching or surpassing sternites II–IV and laterally between IV–VI. abdominal apex. ABDOMEN: Apex rounded; Males are similar to males of T. steineri , T. nam- intersegmental sutures carinulate between steroroka, and T. vazimba , but differs from these nites II–IV and laterally between IV–VI; pygospecies by the characters mentioned. This species phore process subtriangular in caudal view (pl. is most similar to T. brucei , but is distinguished 19I), not surpassing posterior margin of pygoby the smaller width of the synthlipsis relative to phore; BPE shorter than basal plate; endosomal the eye, interocular sulcus anterior to the hind struts not reaching posterior margin of DPS (pl. margin of the eye, larger eye in lateral view, eye 21L), with anterior and posterior ventral pronot reaching ventral head margin, and striated cesses (pl. 22G); area of endosomal struts–DPS posterior pronotal lobe. fusion elongate ovoid (pl. 21L); endosoma medi- DESCRIPTION: MALE: Body length: 13.01 mm ally sclerotized (pl. 21L).

(holotype), 11.49–13.01 mm. COLORATION: ETYMOLOGY: The species epithet is an adjec- Blackish with pale yellow postclypeal medial lon- tive in the nominative case and is named for the gitudinal depression, labial segment IV, scape pale basal part of the femora.

basally, meso- and metasterna, corium basally, DISTRIBUTION: Mahajanga and northern mid and hind coxae, trochanters, forefemur ven- region of Toliara provinces (map 11).

trally and basally, mid and hind femora basally, DISCUSSION: The coloration of the medial tibiae medially, tarsi, anterior half of lateroter- longitudinal depression on the postclypeus is gites, abdominal sternites II–VI medially (except narrowly or broadly pale yellow in some speciat margins of II and III and posteromedial spots mens. The variability of antennal color patterns on III–VI), and anterolateral spots on sternites. could not be fully assessed because parts of the

104 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

antennae are missing in most specimens. In one DESCRIPTION: MALE: Body length: 12.61 specimen, the basal antennal segments are light mm (holotype), 11.00– 12.70 mm. COLORbrown with segments V – VII pale yellow. This ATION: Blackish with pale yellow scape basally, may be a variant as all other specimens have a antennal segments VI and VII, corium basally, dark brown to black scape and pedicel. It is likely trochanters, forefemur ventrally, mid and hind that segments V – VII are always pale yellow. femora basally, tibiae medially, tarsi, anterior HOLOTYPE: Male : Madagascar: Toliara: half of laterotergites, abdominal sternites III – VI Beroboka village , 45 km NE Morondava, medially except at margins, and anterolateral 19.9775 ° S 44.82483 ° E, 131 m, 03 Dec 2008 –11 spots on sternites. VESTITURE: Sparse ; other Dec 2008, M. Irwin, R. Harin’Hala (00045571) features as in generic description. STRUCTURE: ( CAS). HEAD: Subquadrate in lateral view (pl. 13K) ; as PARATYPES: Madagascar: Mahajanga: 160 km N of long as wide in dorsal view ; ventrally with deep Maevatanana on RN 04 , Ampijoroa National Park , medial depression ; anteocular region longer 16.31933 ° S 46.81333 ° E, 43 m, 07 Feb 2005 –19 Feb

than postocular (pl. 13K); postclypeus with 2005, M. Irwin, R. Harin’Hala, 13 (00045511) (CAS).

Toliara: Beroboka village, 45 km NE Morondava, deep, broad medial longitudinal depression to 19.9775 ° S 44.66533 ° E, 128 m, 16 Oct 2008 –24 Oct interocular sulcus; synthlipsis about two times 2008, M. Irwin, R. Harin’Hala, 13 (00045574) (CAS); width of eye; interocular sulcus near hind mar- 09 Nov 2008 – 17 Nov 2008, M. Irwin, R. Harin’Hala, gin of eye (pl. 13B); postocular narrow in dorsal 13

(00044958) (CAS);

17

Nov 2008–25 Nov

2008,

M.

view (pl. 13A); ocelli large, located on shallow Irwin, R. Harin’Hala, 23 (00007161, 00045159)

(USNM); 19 Dec 2008 – 27 Dec 2008, M. Irwin, R. median tubercle (pl. 13K), separated by less than Harin’Hala, 13 (00044937) (UCR). Beroboka village, diameter of ocellus; eye about one-third of head 45 km NE Morondava, 19.9775 ° S 44.82483 ° E, 131 m, length, not reaching dorsal and ventral head 30 Oct 2007 – 06 Nov 2007, M. Irwin, R. Harin’Hala, 23 surfaces; antenna inserted dorsally on head; (00006972, 00006975) (BMNH), 13 (00006977) (CAS); pedicel about one-fourth longer than scape, 22 Nov 2007 – 30 Nov 2007, M. Irwin, R. Harin’Hala,

slightly curved; labium stout; labial segment III 23 (00007073, 00044898) (CAS), 23 (00045223,

00045525) (SU); 17 Nov 2008 – 25 Nov 2008, M. Irwin, ventrally straight (pl. 13K). THORAX: Anterior R. Harin’Hala, 13 (00045070) (CAS); 25 Nov 2008 –03 pronotal lobe with very small anterolateral pro- Dec 2008, M. Irwin, R. Harin’Hala, 13 (00045705) jections (pl. 15B); posterior pronotal lobe (UCR). Unknown: 13 (00045556) (CAS). smooth; pronotal transverse furrow divided by

paramedian longitudinal ridges (pl. 15B); lateral Toxopus brucei , new species depressions of posterior pronotal lobe smooth;

Plates 3, 7, 10; map 12 hemelytron reaching or surpassing abdominal DIAGNOSIS: Males recognized among other apex. ABDOMEN: Apex rounded or sinuate; species in this genus by the coloration, sparse intersegmental sutures carinulate between stervestiture, head as long as wide, smooth posterior nites II–IV and laterally between IV–VI; pygopronotal lobe, and intersegmental sutures cari- phore process subtriangular in caudal view (pl. nulate between sternites II–IV and laterally 19I), not surpassing posterior margin of pygobetween IV–VI. This species is similar to T. stei- phore; BPE shorter than basal plate; endosomal neri, T. namoroka , and T. vazimba , but differs struts not reaching posterior margin of DPS (pl. from these species by the features mentioned. 21L), with anterior and posterior ventral pro- This species is most similar to T. basalis , from cesses (pl. 22G); area of endosomal struts–DPS which it differs by the larger synthlipsis width, fusion elongate ovoid (pl. 21L); endosoma mediinterocular sulcus near the hind margin of the ally sclerotized (pl. 21L).

eye, smaller eye width in lateral view, eye not ETYMOLOGY: This species epithet is a noun in reaching ventral head margin, and smooth pos- the genitive case and is named in memoriam after terior pronotal lobe. the senior author’s uncle, Thomas Allen Bruce.

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 105

DISTRIBUTION: Mahajanga and south-central Toxopus farafangana , new species

region of Toamasina provinces (map 12). Plates 3, 7, 10; map 11 DISCUSSION: The coloration is slightly vari- DIAGNOSIS: Males are recognized among other

able with antennal segment V pale yellow, the Toxopus species by the coloration, ovoid head

corium, except basally, brown, the meso- and shape in lateral view, anteocular region as long as

metasterna and sternite II with a medial yellow postocular, broad postocular, labial segment III

spot, and the medial spots on the sternites are ventrally convex, smooth posterior pronotal lobe,

not be bordered by dark brown margins. and BPE shorter than the basal plate. Males are HOLOTYPE: Male : Madagascar: Mahajanga: similar to T. pallidus , but the dark brown to black-

Besalampy District, Analangidro dry forest, 7 km ish posterior propleural lobe and thoracic sterna,

NE of Besalampy, 16.6915 ° S 44.5235 ° E, 09 Oct predominately dark brown to blackish ventral

2007– 16 Oct 2007, M. Irwin, R. Harin’Hala abdominal surface, interocular sulcus near the

(00007133) (CAS). hind margin of the eye, labial segment III ven- PARATYPES: Madagascar: Mahajanga: Besalampy trally convex, smooth posterior pronotal lobe, and

District, Analangidro dry forest, 7 km NE of BPE shorter than the basal plate distinguish T.

Besalampy, 16.6915 ° S 44.5235 ° E, 61 m, 02 Oct 2007 – farafangana from T. pallidus .

09 Oct 2007, M. Irwin, R. Harin’Hala, 13 (00007039) DESCRIPTION: MALE: Body length: 10.19 mm

(CAS); 09 Oct 2007 – 16 Oct 2007, M. Irwin, R.

Harin’Hala, 13 (00007217) (SU); 14 Dec 2007 –21 (holotype), 9.05–10.66 mm. COLORATION:

Dec 2007, M. Irwin, R. Harin’Hala, 13 (00045338) Blackish with pale yellow anterior margin of

(UCR). Besalampy District, Marofototra dry forest, antennifer, antennal segment V (except basally)

17 km W of Besalampy, 16.72166 ° S 44.42366 ° E, 52 and VI and VII, corium basally, mid and hind

m, 26 Nov 2007 – 03 Dec 2007, M. Irwin, R. trochanters, forefemur ventrally, mid and hind

Harin’Hala, 13 (00044822) (CAS), 13 (00044972) femora basally and subapically, foretibia subbas-

(BMNH); 10 Dec 2007 – 17 Dec 2007, M. Irwin, R.

Harin’Hala, 23 (00007043, 00007156) (AMNH), 23 ally and apically, mid and hind tibiae apically and

(00007160, 00007170) (USNM), 23 (00007182, medially, tarsi, anterior half of laterotergites,

00007223) (CAS), 13 (00045011) (BMNH); 17 Dec small posterior paramedian stripes on abdomi-

2007– 24 Dec 2007, M. Irwin, R. Harin’Hala, 13 nal sternites II–VII, lateral margin of sternites II

(00045155) (SU). Besalampy District, Marofototra and III, and anterolateral margin of sternites IV–

palm forest, 17 km W of Besalampy, 16.71666 ° S

44.41666 ° E, 11 m, 10 Dec 2007 – 17 Dec 2007, M. VII. Labium except basal two-thirds of segment

Irwin, R. Harin’Hala, 13 (00045286) (CAS). Mainti- II, posterior pronotal lobe, and remainder of

rano District Asondrodava dry forest, 15 km N of corium brown. VESTITURE: Dense; other fea-

Maintirano, 17.96533 ° S 44.0355 ° E, 61 m, 03 Dec tures as in generic description. STRUCTURE:

2007– 10 Dec 2007, M. Irwin, R. Harin’Hala, 13 HEAD: Ovoid in lateral view (pl. 13F); longer

(00006140) (CAS); 10 Dec 2007 – 17 Dec 2007, M. than wide in dorsal view; ventrally with shallow

Irwin, R. Harin’Hala, 13 (00007012) (UCR). Main-

tirano District, 50 km E of Maintirano, 18.004 ° S anteromedial depression; anteocular region as

44.452 ° E, 274 m, 25 Oct 2007 – 01 Nov 2007, M. long as postocular (pl. 13J); postclypeus with

Irwin, R. Harin’Hala, 13 (00007193) (CAS). Nam- deep, broad medial longitudinal depression to

oroka village, Befatika Andranovory, 7 km NW middle of interocular area; synthlipsis about 1.5

Vilanandro village, 16.47333 ° S 45.39133 ° E, 122 m, 09 times width of eye; interocular sulcus near hind

Nov 2007– 16 Nov 2007, M. Irwin, R. Harin’Hala, 13

(00007131) (CAS).

Reserve Speciale de Bemarivo, margin of eye (pl. 13B); postocular moderately

23.8 km 223° SW Besalampy, 16.925 ° S 44.36833 ° E, broad in dorsal view (pl. 13B); ocelli large,

30 m, 19 Nov 2002 – 23 Nov 2002, Fisher et al., 13 located on shallow median tubercle (pl. 13K),

(00006149) (CAS). Toamasina: Botanic Garden near separated by less than diameter of ocellus; eye

entrance to Andasibe National Park, 18.92633 ° S about one-third of head length, not reaching

48.40783 ° E, 1025 m, 05 Sep 2001 – 19 Sep 2001, M. dorsal and ventral head surfaces; antenna

Irwin, R. Harin’Hala, 13 (00044971) (CAS). inserted dorsally on head; pedicel about one-

106 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

fourth longer than scape, slightly curved; labium Irwin, F. Parker, R. Harin’Hala, 13 (00045346) (CAS); slender; labial segment III shorter than II, ven- 03 Mar 2007 – 10 Mar 2007, M. Irwin, F. Parker, R. trally convex (pl. 13J). THORAX: Anterior prono- Harin’Hala, 33 (00007173, 00007175, 00007252)

(USNM); 06 Sep 2007 – 13 Sep 2007, M. Irwin, F. Parker, tal lobe with small anterolateral projections (pl. R. Harin’Hala, 13 (00007052) (CAS); 08 Nov 2007 –22 15B); posterior pronotal lobe smooth; pronotal Nov 2007, M. Irwin, F. Parker, R. Harin’Hala, 13 transverse furrow divided by paramedian longi- (00045221) (SU); 20 Dec 2007 – 27 Dec 2007, M. Irwin, tudinal ridges (pl. 15B); lateral depressions of F. Parker, R. Harin’Hala, 23 (00045003, 00045332) posterior pronotal lobe weakly transversely stri- (AMNH); 03 Jan 2008 – 09 Jan 2008, M. Irwin, F. Parker,

R. Harin’Hala, 13 (00044965) (CAS); 09 Jan 2008 –17 ated; hemelytron reaching or surpassing abdomi-

Jan 2008, M. Irwin, F. Parker, R. Harin’Hala, 13 nal apex. ABDOMEN: Apex rounded; (00044938) (UCR); 16 Mar 2008 – 22 Mar 2008, M. intersegmental sutures carinulate between ster- Irwin, F. Parker, R. Harin’Hala, 23 (00006729, nites II and III; pygophore process subtriangular 00044968) (CAS). Manombo Special Reserve camp site, in caudal view (pl. 19I), not surpassing posterior 32 km SSE of Farafangana, 23.02183 ° S 47.72 ° E, 36 m, margin of pygophore, apex rounded; BPE shorter 12 Nov 2004 – 23 Nov 2004, M. Irwin, R. Harin’Hala,

13 (00045531) (CAS); 15 Dec 2004 – 26 Dec 2004, M. than basal plate; endosomal struts reaching pos- Irwin, R. Harin’Hala, 13 (00044890) (SU); 16 Jan terior margin of DPS (pl. 21J, K), with anterior 2005– 20 Jan 2005, M. Irwin, R. Harin’Hala, 13 and posterior ventral processes (pl. 22G); area of (00045258) (UCR).

endosomal struts–DPS fusion ovate; endosoma

medially sclerotized (pl. 21L). Toxopus fisheri , new species

ETYMOLOGY: The species epithet is a noun in Plates 3, 7, 10, 19H, 20K, 21J; map 12 apposition and is named after a locality near DIAGNOSIS: Males are recognized among which known specimens were collected. other species in this genus by the sparse vesti-

DISTRIBUTION: Southeastern region of the ture, coloration, elongate cylindrical head in lat- Fianarantsoa province (map 11). eral view, ventrally flat head surface, sublateral

DISCUSSION: In some specimens, the color- antennal insertion, labial segment III ventrally ation is slightly variable with the posterior pro- straight, and intersegmental sutures carinulate notal lobe dark brown and the abdominal between sternites II and III and laterally between ventrolateral pale yellow spots sometimes larger III–VI. This species is similar to T. italaviana , T. on some sternites or absent. griswoldi , T. antsiranana , and T. toliara , but dif-

HOLOTYPE: Male: Madagascar: Fianarant- fers from these species by the ventrally flat head soa: 50 km S of Farafangana, Mahabo Mananivo, surface. Toxopus fisheri are also similar to T. Ampitavananima Forest, 23.12983 ° S 47.717 ° E, 34 insignis , from which it differs by the dark brown m, 17 Feb 2007 – 24 Feb 2007, M. Irwin, F. Parker, to blackish postclypeal depression, pale yellow R. Harin’Hala (00007034) ( CAS). maxillary plates and antennifers, and interseg-

PARATYPES: Madagascar: Fianarantsoa: 50 km S of mental sutures carinulate between sternites II Farafangana, Mahabo Mananivo, Ampitavananima For- and III and laterally between III – VI.

est, 23.12983 ° S 47.717 ° E, 34 m, 13 Jan 2007 –20 Jan

DESCRIPTION: MALE: Body length: 10.19 mm 2007, M. Irwin, F. Parker, R. Harin’Hala, 23 (00044948,

00044974) (CAS); 20 Jan 2007 – 26 Jan 2007, M. Irwin, (holotype), 9.84–11.09 mm. COLORATION: F. Parker, R. Harin’Hala, 23 (00006147, 00007265) Blackish with pale yellow maxillary plates dor- (CAS); 26 Jan 2007 – 04 Feb 2007, M. Irwin, F. Parker, R. sally, antennifer, scape basally, pedicel basally, Harin’Hala, 23 (00044798, 00045114) (CAS), 23 base of antennal segment III, antennal segment

(00045438, 00045524) (BMNH); 26

Jan 2007–04

Feb

VII, corium basally, coxae, trochanters, forefe- 2007, M. Irwin, R. Harin’Hala, 13 (00045049) (UCR);

04 Feb 2007 – 10 Feb 2007, M. Irwin, F. Parker, R. mur basally and subapically, mid and hind fem- Harin’Hala, 23 (00007080, 00007207) (CAS); 17 Feb ora except medially, foretibia medially except 2007– 24 Feb 2007, M. Irwin, F. Parker, R. Harin’Hala, lateral spots, mid and hind tibiae subbasally and 13 (00007230) (SU); 24 Feb 2007 – 03 Mar 2007, M. medially, tarsi, laterotergites II and VII, anterior

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 107

half of laterotergites III – VI, and abdominal ster- eus, meso- and metasterna, and mid tibia medinites laterally and medially except intersternal ally pale yellow in some specimens. sutures. Antennal segments V, VI, and apex of HOLOTYPE: Male: Madagascar: Antsiranana: VII and labium brown. VESTITURE: Sparse ; Forêt de Binara, 7.5 km 230° SW Daraina, other features as in generic description. STRUC- 13.255 ° S 49.61666 ° E, 375 m, 01 Dec 2003, B.L. TURE: HEAD: Cylindrical in lateral view (pl. Fisher (00045439) ( CAS). 13J) ; longer than wide in dorsal view; ventrally PARATYPES: Madagascar: Antsiranana: Forêt Bekaflat ; anteocular region longer than postocular (pl. raoka, 6.8 km 60° ENE Daraina , 13.00277 ° S 49.01166 ° E,

150 m, 07 Dec 2003, B.L. Fisher, 13 (00045028) (UCR), 13K); postclypeus with shallow, narrow medial

13 (00045100) (SU), 23 (00045143, 00045592) (CAS). longitudinal depression to middle of interocular Forêt d’Ampondrabe, 26.3 km 10 ° NNE Daraina, area (pl. 13B); synthlipsis about two times width 12.97 ° S 49.7 ° E, 175 m, 10 Dec 2003, B.L. Fisher, 13 of eye; interocular sulcus near hind margin of (00045087) (CAS), 23 (00045141, 00045148) (SU), 13 eye (pl. 13B); postocular broad in dorsal view (pl. (00045636) (UCR). Forêt de Binara, 7.5 km 230° SW

Daraina, 13.255 ° S 49.61666 ° E, 375 m, 01 Dec 2003, B.L. 13B); ocelli small, located on shallow median

Fisher, 13 (00045469) (CAS). Marojejy National Park, tubercle (pl. 13J), separated by a distance less 5 km W Manantenina village, Camp Mantella, than or equal to diameter of ocellus; eye about 14.43816 ° S 49.774 ° E, 490 m, 15 Dec 2004 – 20 Dec 2004, one-fourth of head length, not reaching dorsal M. Irwin, R. Harin’Hala, 13 (00006969) (CAS); 05 Jan and ventral head surfaces; antenna inserted sub- 2005– 10 Jan 2005, M. Irwin, R. Harin’Hala, 13 laterally on head; pedicel about two-fifths longer (00006968) (UCR). Rés. Analamerana, 28.4 km 99°

Anivorano-Nord, 12.74666 ° S 49.49472 ° E, 60 m, 05 Dec than scape, slightly curved; labium slender; labial

2004, B.L. Fisher, 13 (00045431) (CAS). segment III ventrally straight (pl. 13K). THORAX: Anterior pronotal lobe with small anterolateral projections (pl. 15B); posterior pronotal lobe Toxopus griswoldi , new species weakly transversely striated; pronotal transverse Plates 3, 4, 7, 10, 23E; map 10 furrow divided by paramedian longitudinal DIAGNOSIS: Males recognized among other ridges (pl. 15B); lateral depressions of posterior Toxopus species by the small body size, sparse pronotal lobe transversely striated; hemelytron vestiture, coloration, elongate cylindrical head in surpassing abdominal apex. ABDOMEN: Apex lateral view, sublateral antennal insertion, labial rounded; intersegmental sutures carinulate segment III ventrally straight, and intersegmenbetween sternites II and III and laterally between tal sutures carinulate between sternites II–IV and III–VI; pygophore process curved in lateral view laterally between IV–VII. In addition to the (pl. 20K), elongate in caudal view (pl. 19H), sur- generic diagnostic characters, the slender, slightly passing dorsal margin of pygophore; BPE shorter curved hind tibia distinguishes females of this than basal plate; endosomal struts reaching pos- species from other known Toxopus females. terior margin of DPS (pl. 21J), with anterior and Males are similar to T. italaviana , T. insignis , T. posterior ventral processes (pl. 22G); area of fisheri , T. antsiranana , and T. toliara , but differs endosomal struts–DPS fusion ovate (pl. 21J); from these species by the small body size. The endosoma with sclerotized, hairlike, denticles on sparse vestiture further distinguishes this species posterior margin (pl. 21J). from T. italaviana , T. antsiranana , and T. toliara , ETYMOLOGY: The species epithet is a noun in while the intersegmental sutures carinulate the genitive case and is named after Brian Fisher, between sternites II–IV and laterally between who collected the majority of specimens exam- IV–VII differentiate T. griswoldi from T. insignis ined for this species. and T. fisheri . DISTRIBUTION: Antsiranana province (map 12). DESCRIPTION: MALE: Body length: 9.28 mm DISCUSSION: The coloration varies with the (holotype), 8.74–9.76 mm. COLORATION: small medial longitudinal stripe on the postclyp- Blackish with pale yellow clypeus; antennifer;

108 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

scape except apically, antennal segment VII FEMALE: Body length: 11.75 mm ; In addition except apically, collar, posterolateral pronotal to characters mentioned in the generic descripmargins, meso- and metasterna medially, scutel- tion: STRUCTURE: HEAD: Synthlipsis about 3.5 lar processes, corium basally, coxae, trochanters, times width of eye. THORAX: Hind tibia slender, forefemur basally and subapically, mid and hind slightly curved. ABDOMEN: Intersegmental femora except medially, tibiae medially, tarsi, lat- sutures carinulate laterally between sternites II – erotergite II, anterior area of laterotergites III – VI ; external genitalia as in plate 23E; bursa copu- VII, and abdominal sternites medially and latrix membranous, damaged; vermiform gland laterally. Antennal segment VI and apex of VII and lateral spermatheca missing. and labium brown. VESTITURE: Very sparse ; ETYMOLOGY: The species epithet is a noun in other features as in generic description. STRUC- the genitive case and is named after Charles TURE: HEAD: Cylindrical in lateral view (pl. Griswold , who collected the only known female 13J) ; longer than wide in dorsal view; ventrally specimen known for this species. with shallow anteromedial depression; anteocu- DISTRIBUTION: Mahajanga, Toliara, and the lar region longer than postocular (pl. 13K) ; northern area of the Antananarivo provinces postclypeus with shallow, narrow medial longi- (map 10). tudinal depression to middle of interocular area DISCUSSION: Some specimens have pale yel- (pl. 13B); synthlipsis about two times width of low mandibular plates. The female specimen was eye; interocular sulcus near hind margin of eye associated with males based on morphology, (pl. 13B); postocular broad in dorsal view (pl. geographic proximity, and molecular data ( COI 13 B); ocelli small, located on shallow median pairwise genetic distance 0.76%). tubercle (pl. 13J), separated by diameter of ocel- HOLOTYPE: Male: Madagascar: Toliara: Cap lus ; eye about one-fourth of head length, not Ste Marie Special Reserve , 74 km S of Tsihombe, reaching dorsal and ventral head surfaces ; 25.58766 ° S 45.163 ° E, 37 m, 23 Nov 2003 –30 Nov antenna inserted sublaterally on head; pedicel 2003, M. Irwin, F. Parker, R. Harin’Hala about one-fourth longer than scape, slightly (00045173) ( CAS). curved ; labium slender; labial segment III ven- PARATYPES: Madagascar: Antananarivo: 46 km trally straight (pl. 13K). THORAX: Anterior pro- NE of Ankazobe: Ambohitantely , 18.198 ° S 47.2815 ° E, notal lobe with small anterolateral projections 700 m, 28 Dec 2003 – 10 Jan 2004, M. Irwin, R. (pl. 15B) ; posterior pronotal lobe transversely Harin’Hala, 13 (00045139) ( CAS) ; 14 Nov 2004 –29 striated; pronotal transverse furrow divided by Nov 2004, M. Irwin, R. Harin’Hala, 13 (00045132) ( UCR). Mahajanga: Ambovomamy Belambo , 20 km paramedian longitudinal ridges (pl. 15B) ; lateral NW of Port Berger , 15.45116 ° S 47.61333 ° E, 33 m, 28 depressions of posterior pronotal lobe trans- Oct 2007– 06 Nov 2007, M. Irwin, F. Parker, R. versely striated ; hemelytron surpassing abdomi- Harin’Hala, 13 (00045234) ( CAS) ; 23 Nov 2007 –08 nal apex. ABDOMEN: Apex rounded ; Dec 2007, M. Irwin, F. Parker, R. Harin’Hala, 13 intersegmental sutures carinulate between ster- (00044962) ( SU) ; 08 Dec 2007 – 11 Dec 2007, M. Irwin, F. Parker, R. Harin’Hala, 13 (00045233) ( AMNH) ; 26 nites II–IV and laterally between IV–VII; pygo- Oct 2008– 01 Nov 2008, M. Irwin, F. Parker, R. phore process curved in lateral view (pl. 20K), Harin’Hala, 23 (00044994, 00045075) ( CAS), 23 elongate in caudal view (pl. 19H), not surpassing (00045386, 00045645) ( USNM) ; 16 Nov 2008 –23 Nov posterior margin of pygophore; BPE shorter than 2008, M. Irwin, F. Parker, R. Harin’Hala , 13 basal plate ; endosomal struts reaching posterior (00045056) (CAS); 23 Nov 2008 – 30 Nov 2008, M. margin of DPS (pl. 21J, K), with anterior and Irwin, F. Parker, R. Harin’Hala, 13 (00045692) ( UCR). Analamanitra Forest , 14 km NE of Misinjo, 16.13333 ° S posterior ventral processes (pl. 22G) ; area of 45.7 ° E, 20 m, 23 Oct 2007 – 30 Oct 2007, M. Irwin, R. endosomal struts–DPS fusion elongate subquad- Harin’Hala, 13 (00045008) ( CAS) ; 30 Oct 2007 –16 rate (pl. 21K); endosoma with sclerotized, hair- Nov 2007, M. Irwin, R. Harin’Hala, 13 (00007148) like, denticles on posterior margin (pl. 21J, K). ( AMNH) ; 16 Nov 2007 – 20 Nov 2007, M. Irwin, R .

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 109

Harin’Hala, 33 (00007096, 00007176, 00045025) Toxopus insignis , new species (MNHN); 20 Nov 2007 – 04 Dec 2007, M. Irwin, R. Plates 4, 7, 10, 14; map 13 Harin’Hala, 23 (00044799, 00044896) (USNM). Anji- DIAGNOSIS: Males are recognized among aabo, 3 km N Baly village, 16.059 ° S 45.27416 ° E, 7 m,

other species in Toxopus by the sparse vestiture, 07 Nov 2007 – 24 Nov 2007, M. Irwin, R. Harin’Hala,

13 (00007214) (CAS). Besalampy District, Analan- coloration, elongate cylindrical head in lateral gidro dry forest, 7 km NE of Besalampy, 16.6915 ° S view, sublateral antennal insertion, straight pedi- 44.5235 ° E, 61 m, 24 Sep 2007 – 02 Oct 2007, M. Irwin, cel, and labial segment III ventrally straight. This R. Harin’Hala, 13 (00044806) (CAS); 18 Nov 2007 –25 species is similar to T. italaviana , T. griswoldi , T. Nov 2007, M. Irwin, R. Harin’Hala, 13 (00044886)

fisheri , T. antsiranana , and T. toliara , from which (SU). Besalampy District, Marofototra dry forest, 17

km W of Besalampy, 16.72166 ° S 44.42366 ° E, 52 m, 19 it differs by the straight pedicel. The ventrally flat Nov 2007– 26 Nov 2007, M. Irwin, R. Harin’Hala, 13 head surface further differentiates T. insignis (00044880) (CAS); 26 Nov 2007 – 03 Dec 2007, M. males from the mentioned species, except for T. Irwin, R. Harin’Hala, 13 (00044885) (SU). Namoroka fisheri , which can be discerned from by the pale 53 km from Soalala 3 km N Vilanandro Village, yellow postclypeal depression, dark brown max- 16.47333 ° S 45.39133 ° E, 122 m, 16 Nov 2007 –23 Nov

illary plates and antennifers, and intersegmental 2007, M. Irwin, R. Harin’Hala, 13 (00007171)

(BMNH), 13 (00007212) (CAS). Namoroka village, sutures carinulate between sternites II–IV and Befatika Andranovory, 7 km NW Vilanandro village, laterally between IV–VI.

16.47333 ° S 45.39133 ° E, 122 m, 02 Nov 2007 –09 Nov DESCRIPTION: MALE: Body length: 10.43 mm 2007, M. Irwin, R. Harin’Hala, 13 (00045002) (CAS). (holotype), 9.73–11.41 mm. COLORATION: Parc National de Namoroka, 16.9 km 317° NW

Blackish with pale yellow postclypeal medial lon- Vilanandro, 16.40666 ° S 45.31 ° E, 100 m, 12 Nov 2002 –

16 Nov 2002, Fisher et al., 13 (00045095) ( CAS), 13 gitudinal depression, interocular sulcus, scape (00045133) (SU) ; 12 Nov 2002 – 16 Nov 2002, Fisher et basally, posterior margin of mesosternum, al., 1♀ (00045042) ( CAS). Parc National de Nam- corium basally, mid and hind coxae anteriorly, oroka, 17.8 km 329° WNW Vilanandro, 16.37666 ° S trochanters, forefemur ventrally and subapically, 45.32666 ° E, 100 m, 08 Nov 2002 – 12 Nov 2002, Fisher mid and hind femora basally and subapically, et al., 13 (00006435) ( CAS). Toliara: Beroboka vil-

fore- and mid tibiae medially, small subbasal and lage, 45 km NE Morondava, 19.9775 ° S 44.66533 ° E,

128 m, 30 Sep 2008 – 08 Oct 2008, M. Irwin, R. subapical band on hind tibia, anterior half of lat- Harin’Hala, 13 (00044841) (CAS); 08 Oct 2008 –16 erotergites, and sternites laterally and medially Oct 2008, M. Irwin, R. Harin’Hala, 13 (00045649) except intersternal suture. Antennal segments VI (AMNH); 09 Nov 2008 – 17 Nov 2008, M. Irwin, R. and VII, labium, and rest of hind tibia brown. Harin’Hala, 23 (00044915, 00045565) (CAS); 11 Dec

VESTITURE: Very sparse; other features as in 2008– 19 Dec 2008, M. Irwin, R. Harin’Hala, 13

(00045695) (UCR). Beroboka village, 45 km NE generic description. STRUCTURE: HEAD: Cylin- Morondava, 19.9775 ° S 44.82483 ° E, 131 m, 14 Oct drical in lateral view (pl. 13J); longer than wide 2007– 18 Oct 2007, M. Irwin, R. Harin’Hala, 13 in dorsal view; ventrally flat; anteocular region (00045110) (CAS); 18 Oct 2007 – 26 Oct 2007, M. longer than postocular (pl. 13K); postclypeus Irwin, R. Harin’Hala, 23 (00044859, 00045096) with shallow, narrow medial longitudinal depres- (BMNH); 17 Nov 2008 – 25 Nov 2008, M. Irwin, R.

sion to middle of interocular area; synthlipsis Harin’Hala, 13 (00044814) (AMNH), 13 (00045041)

(CAS); 05 Feb 2009 – 13 Feb 2009, M. Irwin, R. about two times width of eye; interocular sulcus Harin’Hala, 13 (00045208) (CAS). Cap Ste Marie Spe- near hind margin of eye (pl. 13B); postocular cial Reserve, 74 km S of Tsihombe, 25.58766 ° S broad in dorsal view (pl. 13B); ocelli small, 45.163 ° E, 37 m, 23 Nov 2003 – 30 Nov 2003, M. Irwin, located on shallow median tubercle (pl. 13J),

F. Parker, R. Harin’Hala,

13

(00045379) (CAS). Parc

separated by less than diameter of ocellus; eye National d’Andohahela, Ihazofotsy Parcelle III,

24.83083 ° S 46.53616 ° E, 80 m, 16 Dec 2002 –26 Dec about one-fourth of head length, not reaching 2002, M. Irwin, F. Parker, R. Harin’Hala, 13 dorsal and ventral head surfaces; antenna (00045333) (CAS); 21 Apr 2003 – 28 Apr 2003, M. inserted sublaterally on head; pedicel about one- Irwin, F. Parker, R. Harin’Hala, 13 (00045635) (UCR). third longer than scape, not distinctly curved;

110 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

posteriorly obscured by endosoma; area of endo-

somal struts–DPS fusion short, subquadrate;

endosoma with sclerotized, hairlike, denticles on

posterior margin (pl. 21J, K).

ETYMOLOGY: The species epithet is an adjec-

tive in the nominative case and is named for the

pale interocular sulcus.

DISTRIBUTION: Mahajanga and northern

Antananarivo provinces (map 13).

DISCUSSION: The pale yellow subapical band

on the femora is sometimes absent.

HOLOTYPE: Male : Madagascar: Antanana-

rivo: 46 km NE of Ankazobe: Ambohitantely,

18.198 ° S 47.2815 ° E, 700 m, 14 Nov 2004 –29 Nov

2004, M. Irwin, R. Harin’Hala (00045091) ( CAS).

PARATYPES: Madagascar: Antananarivo: 46 km NE

of Ankazobe: Ambohitantely, 18.198 ° S 47.2815 ° E, 700

m, 31 Oct 2003 – 07 Nov 2003, M. Irwin, R. Harin’Hala,

13 (00045115) ( CAS); 07 Nov 2003 – 15 Nov 2003, M.

Irwin, R. Harin’Hala, 13 (00045662) ( UCR); 15 Oct

2004– 01 Nov 2004, M. Irwin, R. Harin’Hala, 23

(00007098, 00045080) ( CAS), 23 (00045147, 00045265)

(SU), 33 (00045426, 00045594, 00045670) ( AMNH);

01 Nov 2004 – 14 Nov 2004, M. Irwin, R. Harin’Hala,

13 (00045051) ( UCR), 53 (00045057, 00045125,

00045130, 00045140, 00045671) ( CAS), 13 (00045702)

(SU); 14 Nov 2004 – 29 Nov 2004, M. Irwin, R.

Harin’Hala, 33 (00044920, 00045009, 00045065)

( AMNH), 63 (00044920, 00045009, 00045065, MAP 13. Localities of Toxopus insignis , T. namoroka , 00045073, 00045081, 00045082) ( BMNH), 143 T. politus , T. signoretii , and T. simulans . (00045090, 00045098, 00045108, 00045113, 00045116,

00045121, 00045122, 00045135, 00045138, 00045149, labium stout; labial segment III ventrally straight 00045160, 00045162, 00045168, 00045169, 00045664, (pl. 13K). THORAX: Anterior pronotal lobe with 00045665, 00045668) ( CAS), 23 (00045679, 00045690)

(SU), 23 (00045696, 00045697) ( UCR); 29 Nov 2004 – small anterolateral projections (pl. 15B); poste-

07 Dec 2004, M. Irwin, R. Harin’Hala, 23 (00044856, rior pronotal lobe transversely striated; pronotal 00045030) ( UCR), 33 (00045031, 00045040, 00045043) transverse furrow divided by paramedian longi- ( USNM), 133 (00045046, 00045055, 00045058, tudinal ridges (pl. 15B); lateral depressions of 00045060, 00045066, 00045084, 00045103, 00045126, posterior pronotal lobe transversely striated; 00045146, 00045150, 00045151, 00045158, 00045164) hemelytron surpassing abdominal apex. ABDO- ( CAS), 63 (00045165, 00045659, 00045661, 00045666,

00045673, 00045703) ( MNHN), 23 (00045724, MEN: Apex rounded; intersegmental sutures cari- 00045725) (SU) ; 22 Dec 2004 – 06 Jan 2005, M. Irwin , R. nulate between sternites II – IV and laterally Harin’Hala, 33 (00007068, 00045145, 00045587) between IV – VI ; pygophore process curved in ( USNM). Mahajanga: Analamanitra Forest, 14 km NE lateral view (pl. 20K), elongate in caudal view (pl. of Misinjo, 16.13333 ° S 45.7 ° E, 20 m, 16 Nov 2007 –20 19H), not surpassing posterior margin of pygo- Nov 2007, M. Irwin , R. Harin’Hala, 13 (00007229)

( CAS). Parc National Tsingy de Bemaraha , 3.4 km 93 °E phore ; BPE shorter than basal plate; endosomal

Bekopaka, Tombeau Vazimba, 19.14194 ° S 44.82805 ° E, struts reaching posterior margin of DPS (pl. 21J, 50 m, 06 Nov 2001 – 10 Nov 2001, Fisher et al., 13 K), with anterior ventral processes (pl. 22G), (00045054) ( CAS). Réserve forestière Beanka, 50.7 km

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 111

E Maintirano, 17.88027 ° S 44.46888 ° E, 140 m, 28 Oct ger than postocular (pl. 13K); synthlipsis about 2009– 31 Oct 2009, B.L. Fisher et al., 13 (00006910) two times width of eye; interocular sulcus near ( CAS); 13 (00007081) ( UCR).

hind margin of eye; postclypeus with shallow,

broad medial longitudinal depression to middle

Toxopus italaviana , new species

of interocular area; postocular moderately broad

Plates 4, 7, 10, 13B, 15B; map 12

in dorsal view; ocelli large, located on shallow

DIAGNOSIS: Males are recognized among

median tubercle (pl. 13J), separated by a distance

other Toxopus species by the moderately dense

less than or equal to diameter of ocellus; eye

vestiture, elongate cylindrical head shape in lat-

about one-third of head length, not reaching

eral view, broad postclypeal longitudinal depres-

dorsal and ventral head surfaces; antenna

sion, relatively large ocelli, antenna inserted

inserted sublaterally on head; pedicel about two-

sublaterally on the head, and labial segment III

fifths longer than scape, slightly curved; labium

ventrally convex. This species is similar to T. gris-

stout; labial segment III ventrally convex (pl.

woldi, T. insignis , T. fisheri , T. antsiranana , and

13J). THORAX (pl. 15B): Anterior pronotal lobe

T. toliara , but is distinguished by the diagnostic

with small anterolateral projections; posterior

characters mentioned. The slightly larger ocelli

pronotal lobe weakly transversely striated; pro-

further distinguish T. italaviana from the other

notal transverse furrow divided by paramedian

species, except T. toliara . The intersegmental

longitudinal ridges; lateral depressions of poste-

sutures carinulate between sternites II and III

rior pronotal lobe transversely striated; hemely-

and laterally between III–VII also distinguish

tron surpassing abdominal apex. ABDOMEN:

males of T. italaviana from these species, with

Apex rounded; intersegmental sutures carinulate

the exception of T. griswoldi .

between sternites II and III and laterally between

DESCRIPTION: MALE: Body length: 10.58 mm

III–VII; pygophore process curved in lateral

(holotype), 10.25–10.63 mm. COLORATION:

view (pl. 20K), elongate in caudal view (pl. 19H),

Blackish with pale yellow mandibular and maxil-

not surpassing posterior margin of pygophore;

lary plates dorsally, postclypeus except laterally

BPE shorter than basal plate; endosomal struts

above eye, antennifer, scape basally, antennal

reaching posterior margin of DPS (pl. 21J, K),

segment VII basally, labial segment II ventroapi-

with anterior ventral processes (pl. 22G), poste-

cally, labial segment III ventrobasally, anterolat-

riorly obscured by endosoma; area of endosomal

eral projections of pronotum, mesosternum

struts–DPS fusion short, subquadrate; endosoma

anteromedially and posterolaterally, metaster-

with sclerotized, hairlike, denticles on posterior

num medially, corium basally, coxae, trochan-

margin (pl. 21J, K).

ters, forefemur basally and ventrally, mid femur

ETYMOLOGY: The species epithet is a noun in

basally and subapically, hind femur basally and

apposition and is named after the type locality,

apically, foretibia except basally, mid tibia except

Italaviana, Madagascar.

apically, hind tibia except medially and apically,

DISTRIBUTION: Italaviana, near the Antanan-

laterotergite II, anterior half of dorsal lateroter-

arivo-Fianarantsoa border (map 12).

gites III–VII, ventral laterotergites III–VII, and

HOLOTYPE: Male : Madagascar: Fianarant-

abdominal sternites medially and laterally.

soa: Italaviana, 35 km SSE of Antsirabe,

Antennal segment VI and apex of VII, remainder

20.17333 ° S 47.086 ° E, 1360 m, 27 Feb 2005 –13

of labium, and corium brown. VESTITURE:

Mar 2005, M. Irwin, R. Harin’Hala (00006433)

Moderately dense; other features as in generic

( CAS).

description. STRUCTURE: HEAD (pl. 13B):

PARATYPES: Madagascar: Fianarantsoa: Italaviana,

Cylindrical in lateral view (pl. 13J); longer than 35 km SSE of Antsirabe, 20.17333 ° S 47.086 ° E, 1360 m, wide in dorsal view; ventrally with shallow 09 Dec 2002 – 19 Dec 2002, M. Irwin, R. Harin’Hala, 13 anteromedial depression; anteocular region lon- (00006965) ( CAS); 10 Mar 2003 – 20 Mar 2003, M.

112 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

Irwin, R. Harin’Hala, 13 (00006431) ( CAS), 13 view; ventrally with deep medial depression; (00006966) ( UCR); 15 Jul 2003 – 26 Jul 2003, M. Irwin, anteocular region longer than postocular (pl. R. Harin’Hala, 43 (00006438, 00006439, 00006441, 13K); postclypeus with deep, broad medial lon- 00006445) ( CAS), 23 (00006449, 00006450) ( USNM);

gitudinal depression to interocular sulcus; syn-

16 Oct 2003 – 26 Oct 2003, M. Irwin, R. Harin’Hala, 23

(00006961, 00006963) (SU); 22 Jan 2004 – 04 Feb 2004, thlipsis about 1.5 times width of eye; interocular M. Irwin, R. Harin’Hala, 13 (00006962) ( CAS); 24 Mar sulcus posterior to hind margin of eye (pl. 13B); 2004– 08 Apr 2004, M. Irwin, R. Harin’Hala, 13 postocular narrow in dorsal view (pl. 13A); ocelli (00006428) ( CAS), 23 (00006429, 00006430) ( BMNH); large, located on distinct median tubercle (pl. 27 Feb 2005 – 13 Mar 2005, M. Irwin, R. Harin’Hala, 13

13H), separated by less than diameter of ocellus;

(00006432) ( UCR).

eye about one-third of head length, not reaching

dorsal and ventral head surfaces; antenna

Toxopus melobrunneus , new species

inserted dorsally on head; pedicel about one-

Plates 4, 7, 10; map 11

fourth longer than scape, slightly curved; labium

DIAGNOSIS: Males are recognized among

slender; labial segment III ventrally straight (pl.

other species in Toxopus by a combination of

13K). THORAX: Anterior pronotal lobe with very

the following characters: the large body size,

small anterolateral projections (pl. 15B); poste-

coloration, cylindrical head shape in lateral

rior pronotal lobe smooth; pronotal transverse

view, smooth posterior pronotal lobe and lateral

furrow continuous; lateral depressions of poste-

depressions, continuous pronotal transverse

rior pronotal lobe smooth; hemelytron surpass-

furrow, and intersegmental sutures carinulate

ing abdominal apex. ABDOMEN: Apex rounded;

between sternites II and III. Males are similar to

intersegmental sutures carinulate between ster-

males of T. italaviana , T. griswoldi , T. insignis ,

nites II and III; pygophore process subtriangular

T. fisheri , T. ambohitantely , T. antsiranana , and

in caudal view (pl. 19I), not surpassing posterior

T. toliara , but the continuous pronotal trans-

margin of pygophore; BPE as long as basal plate;

verse suture and intersegmental sutures carinu-

endosomal struts reaching posterior margin of

late between sternites II and III distinguish T.

DPS (pl. 21J, K), with anterior and posterior ven-

melobrunneus from these species. The subtrian-

tral processes (pl. 22G); area of endosomal

gular pygophore process and medial endosomal

struts-DPS fusion elongate ovoid (pl. 21L); endo-

sclerotization pattern further differentiate T.

soma medially sclerotized (pl. 21L).

melobrunneus from these species, except from

ETYMOLOGY: The species epithet is an adjec-

T. ambohitantely .

tive in the nominative case and is named for the

DESCRIPTION: MALE: Body length: 14.95 mm

mostly dark brown legs.

(holotype), 13.96–14.95 mm. COLORATION:

DISTRIBUTION: Northern Fianarantsoa and

Blackish with pale yellow area anterior to

southeastern Toliara provinces (map 11).

interocular sulcus, lateral and posterolateral

DISCUSSION: Some specimens have more pale

margins of ocellar tubercle, anterolaterally from

yellow on the ventral femoral and tibial surfaces

interocular sulcus, antennal segments V (except

and laterotergites, and can be reduced on the

basally) and VI and VII, margins and longitudi-

abdomen.

nal depression of metasternum, corium basally,

HOLOTYPE: Male : Madagascar: Fianarant-

femora basally, foretibia ventrally, anterior area

soa: Parc National Ranomafana, radio tower at

of laterotergites III – VII, abdominal sternites II

forest edge, 21.251 ° S 47.40716 ° E, 1130 m, 30 Dec

medially, anterolateral spots on sternites II – VII,

2005– 08 Jan 2006, M. Irwin, R. Harin’Hala

and ventrolateral spots on sternites III – VII.

(00007059) ( CAS).

VESTITURE: Dense; other features as in generic

PARATYPES: Madagascar: Fianarantsoa: Parc

description. STRUCTURE: HEAD: Cylindrical in National Ranomafana, Belle Vue at Talatakely, 21.2665 ° S lateral view (pl. 13J); longer than wide in dorsal 47.42016 ° E, 1020 m, 04 May 2003 – 16 May 2003, M.

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 113

Irwin, R. Harin’Hala, 13 (00007037) (CAS), 13 cally, anterior one-third of laterotergites, and (00007129) (UCR). Parc National Ranomafana, radio lateral margin of abdominal sternites II–VI. tower at forest edge, 21.251 ° S 47.40716 ° E, 1130 m, 07 Mandibular and maxillary plates, labium, May 2002– 14 May 2002, M. Irwin, R. Harin’Hala, 13

remainder of corium, and tarsi brown. VESTI-

(00044926) (CAS), 13 (00044926) (USNM); 25 Jul

2002– 03 Aug 2002, M. Irwin, R. Harin’Hala, 13 TURE: Dense; other features as in generic (00045238) (SU); 06 Jul 2003 – 17 Jul 2003, M. Irwin, R. description. STRUCTURE: HEAD: Ovoid in lat- Harin’Hala, 13 (00045291) (CAS); 12 Feb 2004 –23 Feb eral view (pl. 13F); longer than wide in dorsal 2004, M. Irwin, R. Harin’Hala, 13 (00007009) (CAS); view; ventrally with shallow medial depression; 24 Jun 2004 – 08 Jul 2004, M. Irwin, R. Harin’Hala, 13

anteocular region longer than postocular (pl.

(00045275) (USNM); 12 Jul 2005 – 25 Jul 2005, M.

Irwin, R. Harin’Hala, 13 (00044927) (USNM); 06 Nov 13K); postclypeus with deep, broad medial lon- 2005– 13 Nov 2005, M. Irwin, R. Harin’Hala, 13 gitudinal depression to interocular sulcus; (00007127) (CAS); 07 Sep 2006 – 20 Sep 2006, M. Irwin, synthlipsis about two times width of eye; R. Harin’Hala, 23 (00007028, 00007196) (SU); 11 Nov interocular sulcus near hind margin of eye (pl. 2006– 18 Nov 2006, M. Irwin, R. Harin’Hala, 23 13B); postocular broad in dorsal view (pl. 13B); (00044935, 00044941) (UCR). Toliara: Forêt Ivohibe

ocelli large, located on shallow median tubercle

55.0 km N Tolagnaro, 24.56888 ° S 47.20388 ° E, 200 m,

02 Dec 2006 – 04 Dec 2006, B.L. Fisher et al., 13 (pl. 13J), separated by less than diameter of ocel- (00006728) (CAS). Unknown: 33 (00044925, lus; eye about one-third of head length, not 00044932, 00045243) (CAS). reaching dorsal and ventral head surfaces; antenna inserted slightly sublaterally on head;

Toxopus miandritsara , new species pedicel about one-fourth longer than scape, Plates 4, 7, 11; map 10 slightly curved; labium stout; labial segment III

DIAGNOSIS: Males are recognized among ventrally convex (pl. 13J). THORAX: Anterior other species in this genus by a combination of pronotal lobe with small anterolateral projecthe following characters: the coloration, ovoid tions (pl. 15B); posterior pronotal lobe smooth; head shape in lateral view, anteocular region lon- pronotal transverse furrow divided by parameger than postocular, broad postocular, antenna dian longitudinal ridges (pl. 15B); lateral depresinserted slightly sublaterally on the head, and sions on pronotum smooth; hemelytron reaching smooth posterior pronotal lobe and lateral or surpassing abdominal apex. ABDOMEN: Apex depressions. This species is similar to T. simulans rounded; intersegmental sutures carinulate and T. toamasina , but differs from these species between sternites II and III; pygophore process by the light brown proximal part of the veins on subtriangular in caudal view (pl. 19I), not surthe hemelytra, dark brown ventral abdominal passing posterior margin of pygophore; BPE as surface, larger synthlipsis width, broad postocu- long as basal plate; endosomal struts reaching lar, slight sublateral antennal insertion on the posterior margin of DPS (pl. 21J, K), with antehead, and smooth posterior pronotal lobe and rior and posterior ventral processes (pl. 22G); lateral depressions. Toxopus miandritsara is fur- area of endosomal struts–DPS fusion hexagonal; ther differentiated from T. toamasina by the endosoma medially sclerotized (pl. 21L). intersegmental sutures carinulate between ster- ETYMOLOGY: The species epithet is a noun in nites II and III. apposition and is named after the type locality,

DESCRIPTION: MALE: Body length: 10.06 Miandritsara Forest, Madagascar.

mm (holotype), 9.51–10.06 mm. COLOR- DISTRIBUTION: Miandritsara Forest in the

ATION: Blackish with pale yellow corium Fianarantsoa province (map 10).

basally, proximal wing veins, mid and hind DISCUSSION: The transverse suture between coxae basally, mid and hind trochanters, femora the meso- and metasterna and the ventrolateral basally and ventrally, foretibia subbasally and spots on the anterior abdominal sternites are apically, mid and hind tibiae medially and api- pale yellow in some specimens.

114 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

HOLOTYPE: Male: Madagascar: Fianarantsoa: in lateral view (pl. 13K); longer than wide in dor- Miandritsara Forest , 40 km S Ambositra, 20.79266 ° S sal view; ventrally with deep medial depression; 47.17566 ° E, 825 m, 26 Mar 2007 – 05 Apr 2007, M. anteocular region longer than postocular (pl. Irwin, R. Harin’Hala (00044852) ( CAS). 13K); postclypeus with shallow, narrow medial

PARATYPES: Madagascar: Fianarantsoa: Miandrit- longitudinal depression to interocular sulcus ; sara Forest , 40 km S Ambositra, 20.79266 ° S synthlipsis about two times width of eye ; interoc- 47.17566 ° E, 825 m, 13 Nov 2006 – 22 Nov 2006, M.

ular sulcus near hind margin of eye (pl. 13B); Irwin, R. Harin’Hala, 13 (00044848) ( CAS); 22 Nov

2006– 01 Dec 2006, M. Irwin, R. Harin’Hala, 13 postocular narrow in dorsal view (pl. 13A); ocelli (00044826) ( CAS); 10 Dec 2006 – 19 Dec 2006, M. large, located on shallow median tubercle (pl. Irwin, R. Harin’Hala, 13 (00006148) ( UCR); 28 Dec 13K), separated by less than diameter of ocellus; 2006– 04 Jan 2007, M. Irwin, R. Harin’Hala, 23 eye about one-fourth of head length, not reach- (00044834, 00044970) ( CAS), 13 (00044973) (SU); 31 ing dorsal and ventral head surfaces; antenna Jan 2007 – 11 Feb 2007, M. Irwin, R. Harin’Hala, 13

inserted dorsally on head; pedicel about one- (00006481) ( UCR).

third longer than scape, slightly curved; labium

stout; labial segment III ventrally straight (pl. Toxopus namoroka , new species 13K). THORAX: Anterior pronotal lobe with very

Plates 4, 7, 11; map 13 small anterolateral projections (pl. 15B); poste-

DIAGNOSIS: Males are recognized among other rior pronotal lobe smooth; pronotal transverse Toxopus species by a combination of the following furrow divided by paramedian longitudinal characters: the distinctive dark and pale coloration ridges (pl. 15B); lateral depressions of posterior with red-brownish head and thoracic markings, pronotal lobe smooth; wings spread out in specisparse vestiture, narrow postclypeal depression, mens. ABDOMEN: Apex rounded or sinuate; smooth posterior pronotal lobe, and intersegmen- intersegmental sutures carinulate between stertal sutures carinulate between sternites II–VI and nites II–VI and laterally between VI and VII; laterally between VI and VII. Males are similar to pygophore process subtriangular in caudal view males of T. steineri , T. basalis , T. vazimba , and T. (pl. 19I), not surpassing posterior margin of brucei , but differs from these species by the very pygophore; BPE shorter than basal plate; endodistinctive red-brown head and thorax, narrow somal struts not reaching posterior margin of postclypeal depression, eye about one-fourth of DPS (pl. 21L), with anterior and posterior venthe head length, and intersegmental sutures cari- tral processes (pl. 22G); area of endosomal nulate between sternites II–VI and laterally struts–DPS fusion elongate ovoid (pl. 21L); between VI and VII. endosoma medially sclerotized (pl. 21L).

DESCRIPTION: MALE: Body length: 11.44 mm ETYMOLOGY: The species epithet is a noun in (holotype), 11.44–11.56 mm. COLORATION: apposition and is named after the type locality, Blackish with reddish-brown head, labium, Tsingy de Namoroka National Park, scape, pedicel, antennal segment V, anterior pro- Madagascar.

notal lobe except pronotal longitudinal furrow, DISTRIBUTION: Tsingy de Namoroka National dorsal half of anterior propleural lobe, anterior Park in the Mahajanga province (map 13).

margin of mesopleuron, scutellum, and legs DISCUSSION: The medial pale yellow spots on (except coxae posteriorly and femora basally). the abdominal sternites may be divided by a Antennal segments VI and VII, corium basally, brown medial longitudinal stripe, and the pale anterior half of laterotergites, abdominal ster- yellow spot on sternite VI may be reduced.

nites III – VI medially except at margins, and HOLOTYPE: Male: Madagascar: Mahajanga: anterolateral spots on sternites pale yellow. VES- Parc National de Namoroka , 16.9 km 317° NW

TITURE: Sparse; other features as in generic Vilanandro, 16.40666 ° S 45.31 ° E, 100 m, 12 Nov description. STRUCTURE: HEAD: Subquadrate 2002– 16 Nov 2002, Fisher et al. (00045251) (CAS).

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 115

PARATYPE: Madagascar: Mahajanga: Parc National tudinal depression to interocular sulcus ; synthlip- de Namoroka, 16.9 km 317° NW Vilanandro , sis about 1.5 times width of eye ; interocular sulcus

16.40666

° S

45.31

°

E,

100

m,

12

Nov 2002–16 Nov

2002,

posterior to hind margin of eye (pl. 13B); postocu- Fisher et al., 13 (00006477) (CAS).

lar broad in dorsal view (pl. 13B); ocelli large,

located on shallow median tubercle (pl. 13J), sepa-

Toxopus pallidus , new species rated by less than diameter of ocellus; eye about

Plates 4, 7, 11; map 12 one-third of head length, not reaching dorsal and

DIAGNOSIS: Males are recognized among ventral head surfaces; antenna inserted dorsally other species in Toxopus by the smaller body on head; pedicel about one-fourth longer than size, coloration, ovoid head shape in lateral view, scape, slightly curved; labium stout; labial segment anteocular regions as long as postocular, labial III ventrally straight (pl. 13K). THORAX: Anterior segment III ventrally straight, and BPE as long as pronotal lobe with small anterolateral projections the basal plate. This species is similar to T. fara- (pl. 15B); posterior pronotal lobe transversely strifangana, from which it differs by the the pale yel- ated; pronotal transverse furrow divided by paralow posterior propleural lobe and thoracic sterna, median longitudinal ridges (pl. 15B); lateral predominately pale to pale yellow ventral depressions of posterior pronotal lobe transversely abdominal surface, interocular sulcus posterior striated; hemelytron surpassing abdominal apex. to the hind margin of the eye, labial segment III ABDOMEN: Apex rounded; intersegmental sutures ventrally straight, striated posterior pronotal carinulate between sternites II and III; pygophore lobe, and BPE as long as the basal plate. This spe- process subtriangular in caudal view (pl. 19I), not cies is also similar to T. simulans , but is distin- surpassing posterior margin of pygophore; BPE as guished by the slightly smaller body size, ventral long as basal plate; endosomal struts reaching head depression restricted to the anteocular posterior margin of DPS (pl. 21J, K), ventrally region, anteocular as long as postocular, broader obscured by endosoma; area of endosomal struts– postocular, labial segment III ventrally straight, DPS fusion ovate; endosoma missing.

and intersegmental sutures carinulate between ETYMOLOGY: The species epithet is an adjecsternites II and III. tive in the nominative case and is named for the

DESCRIPTION: MALE: Body length: 9.10 mm pale body coloration.

(holotype), 8.85–9.10 mm. COLORATION: DISTRIBUTION: Ranomafana National Park in Blackish with pale yellow scape basally, antennal the Fianarantsoa province (map 12).

segment V (except basally) and VI and VII; pos- HOLOTYPE: Male : Madagascar: Fianarantterior propleural lobe, ventral half of mesopleu- soa: Parc National de Ranomafana , Vatohararon, metapleuron, thoracic sterna, corium basally, nana River, 4.1 km 231° SW Ranomafana, coxae, trochanters, fore- and mid femora ventrally 21.29 ° S 47.43333 ° E, 1100 m, 27 Mar 2003 –31 and basally, hind femur basally, foretibia ventrally, Mar 2003, Fisher et al. (00045432) ( CAS).

mid and hind tibiae except subbasally and api- PARATYPE: Madagascar: Fianarantsoa: Parc cally, anterior margin of dorsal laterotergites, ven- National Ranomafana, Vohiparara, at broken bridge, tral laterotergites, abdominal sternites except 21.22616 ° S 47.36983 ° E, 1110 m, 08 Apr 2002 –15 Apr

2002, M. Irwin, R. Harin’Hala, 13 (00007192) (CAS). lateral medial spots, and margins of pygophore.

Labium and posterior pronotal lobe brown. VES-

TITURE: Dense; other features as in generic Toxopus parkeri , new species description. STRUCTURE: HEAD: Ovoid in lat- Plates 4, 7, 11, 13K; map 11

eral view (pl. 13F); longer than wide in dorsal DIAGNOSIS: Males are recognized among view; ventrally with shallow anteromedial depres- other species in Toxopus by a combination of the sion; anteocular region as long as postocular (pl. following characters: the coloration, dense vesti- 13J); postclypeus with deep, narrow medial longi- ture, subquadrate head in lateral view that is as

116 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

long as wide in dorsal view, and ventrally on head; pedicel about one-fourth longer than depressed head surface, synthlipsis two times scape, slightly curved; labium stout; labial segment width of an eye, interocular sulcus anterior to the III ventrally straight. THORAX: Anterior pronotal hind margin of the eye, intersegmental sutures lobe with very small anterolateral projections (pl. carinulate between sternites II–VI; and BPE as 15B); posterior pronotal lobe transversely striated; long as basal plate. Males are very similar to T. pronotal transverse furrow divided by paramedian basalis , T. tibialis , and T. brucei , but differ by the longitudinal ridges (pl. 15B); lateral depressions of smaller body size, larger synthlipsis width, and posterior pronotal lobe transversely striated; heminterocular sulcus located anterior to the hind elytron surpassing abdominal apex. ABDOMEN: margin of the eye. The smaller eye size relative to Apex rounded or sinuate; intersegmental sutures the head length, the eyes not reaching the dorsal carinulate between sternites II–VI; pygophore and ventral margins, larger synthlipsis width, process subtriangular in caudal view (pl. 19I), not and intersegmental sutures carinulate between surpassing posterior margin of pygophore; BPE as sternites II–VI separate T. parkeri from T. basa- long as basal plate; endosomal struts not reaching lis. The striated lateral depressions on the poste- posterior margin of DPS (pl. 21L), with anterior rior pronotal lobe and intersegmental sutures and posterior ventral processes (pl. 22G); area of carinulate between sternites II–VI differentiate endosomal struts–DPS fusion elongate ovoid (pl. T. parkeri from T. brucei . 21L); endosoma medially sclerotized (pl. 21L). DESCRIPTION: MALE: Body length: 12.22 mm ETYMOLOGY: The species epithet is a noun in (holotype), 11.52–13.34 mm. COLORATION: the genitive case and is named after Frank Parker, Blackish with pale yellow clypeus, postclypeal who is one of the main CAS collectors for the medial longitudinal depression, bases of scape and taxonomic inventory of Madagascar. pedicel, antennal segment V (except basally) and DISTRIBUTION: Antananarivo, Mahajanga, VI and VII, meso- and metasterna, corium basally, and Toliara provinces (map 11). forecoxa posteriorly, mid and hind coxae except DISCUSSION: The antennae and postclypeus laterally, trochanters, forefemur except dorsally, vary with respect to the extent of the dark brown mid and hind femora basally, foretibia except coloration. basally, mid and hind tibiae medially, tarsi, ante- HOLOTYPE: Male: Madagascar: Toliara: Parc rior half of laterotergites, and abdominal sternites National d’Andohahela, Ihazofotsy Parcelle III, (except pygophore) medially and laterally. Scape 24.83083 ° S 46.53616 ° E, 80 m, 16 Dec 2002 –26 and pedicel medially, antennal segment IV and Dec 2002, M. Irwin, F. Parker, R. Harin’Hala base of V, and postclypeal medial longitudinal (00044875) (CAS). depression brown. VESTITURE: Dense; other PARATYPES: Madagascar: Antananarivo: 46 km NE features as in generic description. STRUCTURE: of Ankazobe: Ambohitantely, 18.198 ° S 47.2815 ° E, 700 HEAD (pl. 13K): Subquadrate in lateral view; as m, 20 Jan 2004 – 01 Feb 2004, M. Irwin, R. Harin’Hala, long as wide in dorsal view; ventrally with shallow 13 (00006973) (UCR), 13 (00006974) (CAS). Mahajanga: Maintirano District Asondrodava dry forest, 15 medial depression; anteocular region longer than km N of Maintirano, 17.96533 ° S 44.0355 ° E, 61 m, 03 postocular; postclypeus with shallow, broad Dec 2007 – 10 Dec 2007, M. Irwin, R. Harin’Hala, 13 medial longitudinal depression to interocular sul- (00007075) (CAS). Toliara: Parc National d’Andohahela, cus; synthlipsis about two times width of eye; Ihazofotsy Parcelle III, 24.83083 ° S 46.53616 ° E, 80 m, 16 interocular sulcus anterior to hind margin of eye; Dec 2002 – 26 Dec 2002, M. Irwin, F. Parker, R. postocular narrow in dorsal view (pl. 13A); ocelli Harin’Hala, 13 (00044884) (CAS); 24 Feb 2003 – 06 Mar 2003, M. Irwin, F. Parker, R. Harin’Hala, 23 large, located on shallow median tubercle, sepa- (00007091, 00044897) (SU), 23 (00045204, 00045249) rated by less than diameter of ocellus; eye about (CAS), 13 (00045381) (UCR); 21 Apr 2003 –28 Apr one-third of head length, not reaching dorsal and 2003, M. Irwin, F. Parker, R. Harin’Hala, 23 (00045052, ventral head surfaces; antenna inserted dorsally 00045377) (USNM). Unknown: 13 (00006976) (CAS).

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