Gibbosella andasibe, 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-

Gibbosella andasibe , new species

Plates 1, 5, 8; map 2

DIAGNOSIS: Males recognized among other species in this genus by the very small body size, coloration, anteocular region shorter than the postocular, postclypeus not depressed, synthlipsis width about 2.5 times the width of an eye, and fossula spongiosa absent on mid tibia. This species is similar to G. megafrons , from which it differs by the light brown anterior pronotal lobe, brown posterolateral margin of pronotum, pale posterior half of scutellum, the flat postclypeus, smaller synthlipsis width, lack of a fossula spongiosa on the mid tibia, and intersegmental sutures carinulate between sternites II–IV.

DESCRIPTION: MALE: Body length: 3.64 mm (holotype). COLORATION: Dark brown with antennal segments VII and VIII, posterolateral margin of pronotum, pleura dorsad of coxae, spot on dorsoposterior margin of mesopleuron, posterior half of scutellum, corium basally, coxae, trochanters, femora basally, laterotergites, except posterior half of VII, and sternites medially and anterolaterally pale. Wing membrane, remainder of antenna, legs, and corium brown. Labium, neck dorsally, and anterior pronotal lobe light brown. VESTITURE: Sparse; other features as in MAP 2. Localities of Gibbosella andasibe , G. elongata , generic description. STRUCTURE: HEAD: Ovoid G. fulva , G. nitida , G. notoconica and G. quadocris . in lateral view (pl. 13F); longer than wide in dorsal view; anteocular region shorter than postocu- – Head shorter, ovate (pl. 13F); anterior pronotal lar; gula flat, conforming to rounded shape of lobe slightly to moderately elevated dorsally head (pl. 13C); postclypeus flat (pl. 13A); synin lateral view; body length <8 mm ...... 3 thlipsis about 2.5 times width of eye; interocular 2. Scutellar processes convergent; hind tibia with sulcus posterior to hind margin of eye; ocelli dark apical half......... mirabilis Chłond small, located on shallow median tubercle (pl. – Scutellar processes not convergent; hind tibia 13I); eye about one-fourth of head length, not yellowish with dark annulus in basal part.. reaching dorsal and ventral head surfaces;........................ elongata Chłond antenna inserted dorsally on head; pedicel as 3. Synthlipsis width three times width of eye; long as scape; antenna 8-segmented; labium generally brown to dark brown with scutel- stout; labial segment III subequal to II, ventrally lar apical projections whitish to pale...... convex (pl. 13C). THORAX: Anterior pronotal................ planiscutum , new species lobe with very small anterolateral projections (pl. – Synthlipsis width two times width of eye; gen- 15B); pronotal transverse furrow continuous; laterally brown to dark brown with scutellar eral depressions on posterior pronotal lobe apical projections brown to dark brown... smooth; scutellar apical projections slightly dor-.................. brunalvus , new species sally directed (pl. 15C); meso- and metasterna

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 39

separated by distinct transverse suture (pl. 15F); mesosternum with medial and paramedial longitudinal depressions; metasternum slightly medially longitudinally depressed posteriorly; legs slender; fossula spongiosa absent on mid tibia; tarsomeres I and II combined subequal to III; hemelytron surpassing abdominal apex; distal part of R absent. ABDOMEN: Apex medially notched; sternites medially convex, intersegmental sutures carinulate between II – IV; pygophore process subtriangular in lateral (pl. 20A) and caudal (pl. 19G, I) views, apex rounded; BPE shorter than basal plate; endosomal struts reaching posterior margin of DPS, with posterior ventral process (pl. 22G); area of endosomal struts–DPS fusion elongate subquadrate. ETYMOLOGY: The species epithet is a noun in apposition and is named after the type locality, Andasibe National Park , Madagascar. DISTRIBUTION: The single known specimen was collected southeast of Andasibe National Park headquarters in the Toamasina province (map 2). HOLOTYPE: Male: Madagascar: Toamasina: 7 km SE of Andasibe National Park headquarters, 18.96266 ° S 48.45266 ° E, 1050 m, 07 Jan 2001 – 22 Jan 2001, M. Irwin, R. Harin’Hala (00044816) ( CAS).

MAP 3. Localities of Gibbosella betampona , G. man-

Gibbosella betampona , new species tella, G. planiscutum , and G. vangocris .

Plates 1, 5, 8; map 3 DIAGNOSIS: Males recognized among other DESCRIPTION: MALE: Body length: 6.40 mm species in Gibbosella by the circular head shape (holotype). COLORATION: Dark brown with in lateral view that is as long as it is wide in dor- pale labial segment IV, antennal segment V apisal view, large ocelli, eyes reaching dorsal and cally, antennal segments VI–VIII, lateral spots ventral head margin, labial segment II longer on meso- and metasterna, coxae, trochanters, than III, mesosternum with large medial depres- forefemur basally, mid and hind femora basally sion, absence of the fossula spongiosa on the mid and subapically, foretibia except medially, mid tibia, and BPE shorter than the basal plate. This and hind tibiae, tarsi, anterior half of lateroterspecies is similar to G. mantella , but the pale pos- gites, paramedian spots on sternites III–V, and terior margin of the pronotum and lateral spots lateral margin of sternite VII. Remainder of on the meso- and metasterna, the brown corium, labium, corium, and basal wing veins brown. larger synthlipsis width, meso- and metasterna VESTITURE: Sparse; other features as in not completely separated by a distinct suture, generic description. STRUCTURE: HEAD: absence of the fossula spongiosa on the mid Ovoid in lateral view (pl. 13C); as long as wide tibia, and shorter BPE relative to the basal plate in dorsal view; anteocular region as long as distinguish this species from G. mantella . postocular (pl. 13C); gula flat, conforming to

40 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

rounded shape of head (pl. 13C); postclypeus PARATYPE: Madagascar: Toamasina: Reserve with shallow, narrow medial longitudinal Betampona, Camp Vohitsivalana , 37.1 km 338 Toamadepression to middle of interocular area (pl. sina, 17.88666 ° S 49.2025 ° E, 520 m, 01 Dec 2005 –03

Dec 2005, Fisher et al., 13 (00007255) (CAS). 13B); synthlipsis about width of eye; interocular sulcus near hind margin of eye (pl. 13B); ocelli large, located on distinct median tubercle (pl. Gibbosella brunalvus , new species 13C); eye about half of head length, not reach- Plates 1, 5, 8, 23A; map 1 ing dorsal head surface, reaching ventral head DIAGNOSIS: Males are recognized among othsurface; antenna inserted dorsally on head; ers in Gibbosella by a combination of the followpedicel about as long as scape; antenna 8-seg- ing characters: the coloration, dense vestiture, mented; labium stout; labial segment III shorter postclypeus not depressed, pronotal transverse than II, ventrally convex (pl. 13C). THORAX: suture divided by paramedian ridges, and scutel- Anterior pronotal lobe with very small antero- lar processes dorsally directed. Females are reclateral projections (pl. 15B); pronotal transverse ognized by the coloration and synthlipsis about furrow continuous; lateral depressions on pos- two times the width of an eye. Males are very terior pronotal lobe smooth; scutellar apical similar to G. fulva , from which G. brunalvus difprojections dorsally directed (pl. 15C); meso- fers by the more abundant pale markings on the and metasterna not completely or weakly sepa- pronotum, scutellum, corium, and sternites, rated by distinct transverse suture (pl. 15E); smaller body size, dense vestiture, head longer mesosternum with large medial depression; than wide, flat postclypeus, and divided pronotal metasternum slightly medially longitudinally transverse furrow, among several other features. depressed; forefemur slightly incrassate; fossula Females are similar to G. planiscutum based on spongiosa absent on mid tibia; tarsomeres I and the features mention in the generic description, II combined shorter than III; hemelytron sur- but are distinguished by the dark brown antennal passing abdominal apex; distal part of R present segment VIII and scutellum, femora basally and but not forming R+M cell (pl. 17B, C). ABDO- tibiae apically and basally pale to light brown, MEN: Apex medially notched; sternites flat, and smaller synthlipsis width. intersegmental sutures carinulate between II– DESCRIPTION: MALE: Body length: 4.89 mm IV and laterally between IV–VI; pygophore pro- (holotype), 4.79–5.13 mm. COLORATION: cess subtriangular in lateral (pl. 20A) and Dark brown with pale color on posterolateral caudal (pl. 19G, I) views, apex rounded; BPE margin of pronotum, scutellar processes, basal shorter than basal plate; endosomal struts half of corium, coxae ventrally, trochanters, femreaching posterior margin of DPS, with poste- ora and tibiae basally, tarsi, anterior half of latrior ventral process (pl. 22G); area of endo- erotergites, and sternites anterolaterally pale. somal struts–DPS fusion ovate (pl. 21A). Antennal segment V apically and segments VI–

ETYMOLOGY: The species epithet is a noun in VIII white. Dorsal head surface, except synthlipapposition and is named after the locality in sis and ocellar tubercle, markings throughout which the two known specimens were collected, anterior pronotal lobe, labium, and antenna Reserve Betampona, Madagascar. brown. VESTITURE: Dense; other features as in

DISTRIBUTION: Betampona Reserve in the generic description. STRUCTURE: HEAD: Ovoid Toamasina province (map 3). in lateral view (pl. 13F); longer than wide in dor-

HOLOTYPE: Male: Madagascar: Toamasina: sal view; anteocular region as long as postocular Reserve Betampona, Camp Vohitsivalana , 37.1 (pl. 13C); gula moderately swollen ventrolaterkm 338 Toamasina , 17.88666 ° S 49.2025 ° E, 520 ally, not distinctly produced beyond ventral head m, 01 Dec 2005 – 03 Dec 2005, Fisher et al. margin (pl. 13E, G–K); postclypeus flat (pl. 13A); (00006143) ( CAS). synthlipsis about two times width of eye; interoc-

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 41

ular sulcus near hind margin of eye (pl. 13B); forefemur with small ventral tubercles (pl. 16D). ocelli small, located on shallow median tubercle ABDOMEN: Intersegmental sutures carinulate (pl. 13I); eye about one-fourth of head length, between sternites II – VI; external genitalia as in not reaching dorsal and ventral head surfaces; plate 23A; bursa copulatrix membranous, with antenna inserted sublaterally on head; pedicel lateral lobes (pl. 23F); vermiform gland and latabout as long as scape; antenna 8-segmented; eral spermathecae damaged. labium stout; labial segment III shorter than II, ETYMOLOGY: The species epithet is a noun in ventrally convex. THORAX: Anterior pronotal the nominative case and is named after the nearly lobe with very small anterolateral projections (pl. uniform dark brown venter. 15B); pronotal transverse furrow divided by DISTRIBUTION: Tsingy de Bemaraha National paramedian longitudinal ridges (pl. 15B) ; lateral Park in Mahajanga province (map 1). depressions on posterior pronotal lobe smooth; DISCUSSION: The coloration is entirely dark scutellar apical projections dorsally directed (pl. brown on the head and anterior pronotal lobe, pale 15C); meso- and metasterna separated by dis- to brown medial stripes on the abdominal sternites tinct transverse suture (pl. 15F); mesosternum in some male specimens. A female specimen was with medial and paramedial longitudinal depres- associated with males based on morphology and sions; metasternum medially longitudinally the same collection event as one other male. depressed; forefemur incrassate; fossula spongi- HOLOTYPE: Male : Madagascar: Mahajanga: osa on mid tibia ; tarsomeres I and II combined Parc National Tsingy de Bemaraha , 3.4 km 93 °E subequal to III ; hemelytron reaching or surpass- Bekopaka, Tombeau Vazimba , 19.14194 ° S ing abdominal apex ; distal part of R present but 44.82805 ° E, 50 m, 06 Nov 2001 – 10 Nov 2001, not forming R +M cell (pl. 17B, C). ABDOMEN: Fisher et al. (00006317) ( CAS). Apex rounded ; sternites convex, shallowly PARATYPES: Madagascar: Mahajanga: Parc National depressed between paramedian ridges on ster- Tsingy de Bemaraha, 3.4 km 93 ° E Bekopaka, Tombeau Vazimba , 19.14194 ° S 44.82805 ° E, 50 m, 06 Nov 2001 – nites II – V, intersegmental sutures carinulate 10 Nov 2001, Fisher et al., 33 (00006122, 00006318, between II – IV and laterally between IV – VI; 00099043) ( UCR), 33 (00006319, 00006321, 00099044) pygophore process subtriangular in lateral (pl. ( SU), 53 (00006322–00006326) ( CAS), 23 (00006327, 20A) and caudal (pl. 19G, I) views, apex rounded; 00006328) ( BMNH), 23 (00006329, 00006455) BPE shorter than basal plate ; endosomal struts ( USNM). Parc National Tsingy de Bemaraha, 10.6 km ESE 123° Antsalova , 19.70944 ° S 44.71806 ° E, 150 m, 16 reaching posterior margin of DPS, with posterior Nov 2001– 20 Nov 2001, Fisher et al., 13 (00045686), ventral process (pl. 22G), fused anterior to fusion 1♀ (00048066) ( CAS). Unknown: 23 (00007154, with DPS; area of endosomal struts–DPS fusion 00044944) ( CAS). circular. FEMALE: Body length: 4.54 mm. Similar to males, but differ in the following characters: COLORATION: Dark brown with pale to Gibbosella conisimilis , new species light brown scape, pedicel, antennal segments Plates 1, 5, 8, 15C, 18A ; map 1 III – IV and basal half of V, labium, femora except DIAGNOSIS: Males are recognized among basal, and tibiae except apical and basal parts. other species in Gibbosella by the larger body Remaining antennal segments, coxae, trochan- size, synthlipsis width 1.5 times the width of an ters, femora basally, tibiae apically and basally, eye, labial segment III ventrally straight, coni- and tarsi whitish to pale brown. STRUCTURE: cally elevated anterior pronotal paramedian In addition to characters mentioned in the lobes in lateral view, anterior pronotal lobe withgeneric description: HEAD: Synthlipsis about two out anterolateral protuberances, continuous protimes width of eye. THORAX: Pronotal longitudi- notal transverse suture, and mesosternum with nal furrow restricted to anterior lobe; lateral large medial depression. This species is similar to depressions on posterior pronotal lobe obsolete; G. notoconica , but the slightly smaller body size,

42 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

pale apex of antennal segment V, dark brown without distinct anterolateral projections; pronopronotum (except anterior and posterolateral tal transverse furrow continuous; lateral depresmargins and disc) and mesosternum (except sions on posterior pronotal lobe smooth; anterolaterally and posteriorly), larger synthlipsis scutellar apical projections dorsally directed; width, interocular sulcus posterior to the hind meso- and metasterna not completely or weakly margin of the eye, smaller ocelli, eye size relative separated by distinct transverse suture (pl. 15E); to the head length and height, and intersegmen- mesosternum with large medial depression; tal sutures carinulate between II and III and lat- metasternum slightly medially longitudinally erally between III – VI distinguish this species depressed; legs slender; fossula spongiosa on mid from G. notoconica . The conical anterior prono- tibia ; tarsomeres I and II combined subequal to tal paramedian lobes are also not as distinct as in III; hemelytron surpassing abdominal apex; dis- G. notoconica . tal part of R present but not forming R +M cell DESCRIPTION: MALE: Body length: 7.92 mm (pl. 17B, C). ABDOMEN (pl. 18A): Apex medially (holotype), 7.86–8.44 mm. COLORATION: notched or sinuate ; sternites medially convex, Dark brown with pale dorsal head surface intersegmental sutures carinulate between II and (except clypeal apex, mandibular plates, anten- III and laterally between III – VI; pygophore pronifers, anterior of interocular sulcus, interocular cess subtriangular in lateral (pl. 20A) and caudal sulcus, and dorsal surface of ocellar tubercle), (pl. 19G, I) views, apex rounded ; BPE as long as scape basally, antennal segment V apically, and basal plate; endosomal struts reaching posterior segments VI – VIII, anterior and posterolateral margin of DPS, ventrally obscured by semiexmargins of pronotum, pronotal disc, lateral lon- tended endosoma; area of endosomal struts–DPS gitudinal depressions of posterior pronotal lobe, fusion ovate (pl. 21A). stridulatory groove, mesosternum anterolaterally ETYMOLOGY: The species epithet is an adjec- and posteriorly, metasternum, scutellum medi- tive in the nominative case and is named for the ally, corium basally, coxae, trochanters, femora similar pronotal appearance of the pronotum in basally and ventrally, mid and hind tibiae medi- G. notoconica . ally, tarsi, anterior margin of laterotergites, and DISTRIBUTION: Marojejy Nature Reserve in sternites medially and anterolaterally. VESTI- Antsiranana province (map 1). TURE: Very sparse ; other features as in generic DISCUSSION: Coloration is variable with the description. STRUCTURE: HEAD: Ovoid in lat- dark brown area around the ocellar tubercle and eral view (pl. 13F) ; longer than wide in dorsal interocular sulcus variable in size. The anterior proview; anteocular region as long as postocular (pl. notal lobe may be dorsally pale, and the pronotal 13C); gula flat, conforming to rounded shape of transverse sulcus entirely or nearly entirely pale. head (pl. 13C); postclypeus with shallow, broad HOLOTYPE: Male: Madagascar: Antsiranana: medial longitudinal depression to middle of Marojejy National Park , 5 km W Manantenina interocular area (pl. 13B) ; synthlipsis about 1.5 village, Camp Mantella, 14.43816 ° S 49.774 ° E, times width of eye; interocular sulcus posterior 490 m, 26 Sep 2005 – 04 Oct 2005, M. Irwin, R. to hind margin of eye ; ocelli small, located on Harin’Hala (00044850) ( CAS). shallow median tubercle (pl. 13I) ; eye about one- PARATYPES: Madagascar: Antsiranana: Marojejy fourth of head length, not reaching dorsal and National Park, 5 km W Manantenina village, Camp ventral head surfaces ; antenna inserted sublater- Mantella, 14.43816 ° S 49.774 ° E, 490 m, 15 Dec 2004 –20 ally on head; pedicel about one-third longer than Dec 2004, M. Irwin, R. Harin’Hala, 13 (00007113) ( CAS), 13 (00045329) ( SU) ; 25 Dec 2004 – 30 Dec 2004, scape; antenna 8-segmented; labium slender; M. Irwin, R. Harin’Hala, 13 (00007260) ( UCR) ; 10 Jan labial segment III subequal to II, ventrally 2005– 15 Jan 2005, M. Irwin, R. Harin’Hala , 13 straight (pl. 13G, H, K). THORAX (pl. 15C): Ante- (00045259) ( CAS) ; 25 Feb 2005 – 04 Mar 2005, M. rior pronotal lobe slightly conical in lateral view, Irwin, R. Harin’Hala , 13 (00007066) ( CAS) ; 25 Mar

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 43

2005– 04 Apr 2005, M. Irwin, R. Harin’Hala, 13 area; synthlipsis about width of eye; interocular (00045427) ( CAS); 04 Apr 2005 – 16 Apr 2005, M. Irwin, sulcus near hind margin of eye (pl. 13B); ocelli R. Harin’Hala, 23 (00007079, 00007083) ( CAS), 23 large sized, located on distinct median tubercle; (00007150, 00007199) ( BMNH); 18 May 2005 –30 May

eye about one-third of head length, reaching dor-

2005, M. Irwin, R. Harin’Hala, 13 (00044819) ( CAS);

30 May 2005 – 11 Jun 2005, M. Irwin, R. Harin’Hala , 13 sal and ventral head surfaces ; antenna inserted (00045317) (SU); 11 Jun 2005 – 28 Jun 2005, M. Irwin , sublaterally on head ; pedicel slightly longer than R. Harin’Hala, 13 (00045495) ( UCR) ; 14 Oct 2005 –22 scape; labium stout; labial segment III shorter Oct 2005, M. Irwin, R. Harin’Hala, 13 (00007054) than II, ventrally convex. THORAX: Anterior pro- ( AMNH). R. N.I. de Marojejy, 8.0 km NW Manan-

notal lobe gibbous (pl. 5) with small anterolateral

tenina, 14.43667 ° S 49.775 ° E, 450 m, 05 Oct 1996 –13

Oct 1996, E. Quinter and T. Nguyen, 13 (00078363) projections; pronotal transverse furrow continu- ( AMNH). Unknown: 13 (00007064) ( CAS). ous; lateral depressions on posterior pronotal lobe striated transversely; scutellar apical projec-

Gibbosella elongata Chłond, 2010 tions dorsally directed (pl. 15C); meso- and

Plates 1, 5, 12; map 2 metasterna separated by distinct transverse suture

(pl. 15F); forefemur incrassate; fossula spongiosa

on mid tibia; tarsomeres I and II combined same

Gibbosella elongata Chłond, 2010, 2522: 64 .

length as III; hemelytron surpassing abdominal

REVISED DIAGNOSIS: Males recognized among apex; distal part of R present but not forming other species in this genus by the larger body R +M cell (pl. 17B, C). ABDOMEN: Sternites medisize, synthlipsis width same as width of an eye, ally convex, intersegmental sutures carinulate labial segment III ventrally convex, elevated between II – IV; pygophore process subtriangular anterior pronotal paramedian lobes in lateral in lateral (pl. 20A) and caudal (pl. 19G, I) views, view, anterior pronotal lobe with small anterolat- apex rounded; BPE as long as basal plate; area of eral projections. Females are recognized by the endosomal struts–DPS subquadrate. FEMALE: cylindrical head, scape surpassing apex of the Body length: 9.40 mm (holotype). Similar to head, and very gibbous anterior pronotal lobe males, but differ in the following characters: with small anterolateral projections. This species COLORATION: Shining, dark brown with pale can be easily distinguished from G. mirabilis by labrum, DFLA, small spots on pronotum and the enlarged, elongated, and much more elevated thorax, coxae, trochanters, basal half of forefemur anterior pronotal lobe and lack of a depression in and two-thirds of mid and hind femora, as well the posterior part of anterior pronotal lobe. as apical part of all femora, tibiae and anterior

REDESCRIPTION: MALE: Body length: 9.90 part of laterotergites. STRUCTURE: In addition mm. COLORATION: Dark brown with pale to characters mentioned in the generic descripbasal part of scape, coxa, trochanters, basal part tion: HEAD (damaged): Slightly convex gula. of forefemur, basal and apical part of mid and THORAX: Anterior pronotal lobe gibbous with hind femur, apical and middle part of mid tibiae, small anterolateral projections (pl. 12); visible hind tibiae except small subapical ring, anterior medial longitudinal sutures on both pronotal part of laterotergites III–VII and middle part of lobes; lateral depressions on posterior pronotal abdominal tergites III–VII. VESTITURE: Sparse; lobe striated transversely. ABDOMEN: Tergites latother features as in generic description. STRUC- erally and laterotergites striated transversely; ter-

TURE: HEAD: Elongate ovoid to cylindrical in gites posterior distinctly rounded, except tergite lateral view; longer than wide in dorsal view; III; sternites with distinct visible fold near lateral anteocular region longer than postocular; gula margin; intersegmental sutures carinulate flat, conforming to rounded shape of head (pl. between sternites II–VII; valvifer I triangular 13C); postclypeus with deep, narrow medial lon- with flat dorsal part and one long setae; valvula I gitudinal depression to middle of interocular small with long hairs placed on small fold visible

44 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

in middle part; valvula III elongate, convolute in gin, labial segment II longer than III, anterior apical part with distinct dome-shaped elevation. pronotal lobe without anterolateral projections, DISTRIBUTION: The type specimen was col- mesosternum with large medial depression, and lected in the Anjanaharibe-Sud Reserve, west of fossula spongiosa on the mid tibia. This species Andapa, which is located in the southern part of is similar to G. brunalvus , but can be distin- Antsiranana province (map 2). The only known guished by the dark brown pronotum, scutelmale specimen was collected in Maroantsetra lum, and corium, larger body size, sparse near the Toamasina-Antsiranana border, which vestiture, head about as long as wide, depressed is in close proximity to the type locality. postclypeus, and pronotal transverse furrow DISCUSSION: Chłond (2010a) recently continuous, among several other features. described this species from a single female repre- DESCRIPTION: MALE: Body length: 7.40 mm sentative. An undetermined male specimen was (holotype), 7.37–7.40 mm. COLORATION: observed to have morphological similarities to G. Dark brown with labial segments III and IV, elongata and G. mirabilis : elongate, cylindrical antennal segments V–VIII, and anterior part of head shape; dorsally protruding anterior pronotal laterotergites pale. VESTITURE: Sparse; other lobe; dorsally oriented scutellar processes; and features as in generic description. STRUCTURE: general dark brown to blackish and pale color- HEAD: Ovoid in lateral view (pl. 13F); about as ation. Although extreme sexual dimorphism long as wide in dorsal view; anteocular region as exists in Gibbosella , we assign the male to G. elon- long as postocular (pl. 13E); gula flat, conformgata based on the small basal dark brown band on ing to rounded shape of head (pl. 13E); postclypthe hind tibia and broader pale bands on the dor- eus with deep, narrow medial longitudinal sal laterotergites; in G. mirabilis , the hind tibia is depression to middle of interocular area; syndark brown on the apical one-third to one-half thlipsis about two times width of eye; interocular and the dorsal laterotergites have more narrow sulcus near hind margin of eye (pl. 13B); ocelli pale bands. Furthermore, the male was collected medium sized, located on shallow median tubernear the border of Toamasina and Antsiranana cle (pl. 13I); eye about one-third of head length, provinces, in close proximity to the type locality almost reaching dorsal and ventral head surfaces; of G. elongata ; G. mirabilis has been collected antenna inserted sublaterally on head; pedicel as from the southwestern region of Toamasina, near long as scape; antenna 8-segmented; labium the Antananarivo-Toamasina border. stout; labial segment III shorter than II, ventrally HOLOTYPE: Female: Madagascar: Antsira- convex (pl. 13E). THORAX: Anterior pronotal nana: Andapa District, Anjanaharibe-Sud lobe without anterolateral projections; pronotal Reserve, 14.69889 ° S 49.45483 ° E, 1600 m, 1960, P. transverse furrow continuous; lateral depressions Soga (MNHN). on posterior pronotal lobe smooth; scutellar api- OTHER MATERIAL EXAMINED: Madagascar: Toama- cal projections slightly dorsally directed (pl. sina: Maroantsetra/Ambodivoangy, 15.43231 ° S 15C); ventrally obscured by card mount; forefe- 49.74013 ° E, 9 m, no date provided, 13 (MNHN).

mur incrassate; fossula spongiosa on mid tibia;

tarsomeres I and II combined shorter than III;

Gibbosella fulva , new species hemelytron surpassing abdominal apex; distal

Plates 1, 5; map 2 part of R present but not forming R+M cell (pl. DIAGNOSIS: Males are recognized among 17B, C). ABDOMEN: Apex medially notched, sterother species in this genus by the shiny brown nites medially obscured by card mount, but color of the body, the head about as long as intersegmental sutures carinulate at least laterally wide in dorsal view, synthlipsis two times the between II–VI; pygophore process subtriangular width of an eye, medium sized ocelli, eyes in lateral (pl. 20A) and caudal (pl. 19G, I) views, almost reaching dorsal and ventral head mar- apex rounded; BPE as long as basal plate; area of

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 45

endosomal struts–DPS fusion basally narrow but view; as long as wide in dorsal view; anteocular distally wider and ovate. region as long as postocular; gula flat, conform- ETYMOLOGY: The species epithet is an adjec- ing to rounded shape of head ; postclypeus with tive in the nominative case and is named after deep, narrow medial longitudinal depression to the shiny brown color of the body. middle of interocular area; synthlipsis less than DISTRIBUTION: Antanambe and Seranambe in width of eye ; interocular sulcus near hind marthe Mananara District of Toamasina (map 2). gin of eye (pl. 13B) ; ocelli large, located on dis- HOLOTYPE: Male : Madagascar: Toama- tinct median tubercle ; eye about half of head sina: Mananara-Nord District, Seranambe , length, reaching dorsal and ventral head sur- 16.24683 ° S 49.83916 ° E, 16 m, Jul 1965, J. Vadon faces ; antenna inserted sublaterally on head; and A. Peyrieras ( MNHN). pedicel as long as scape ; antenna 8-segmented; PARATYPE: Madagascar: Toamasina: Mananara- labium stout ; labial segment III shorter than II, Nord District, Antanambe , 16.43333 ° S 49.85 ° E, 8 m, no ventrally convex. THORAX: Anterior pronotal date provided, J. Vadon and A. Peyrieras, 13 ( MNHN) .

lobe with small anterolateral projections (pl.

15B); pronotal transverse furrow continuous; lat-

Gibbosella mantella , new species eral depressions on posterior pronotal lobe

Plates 1, 5, 8, 13C; map 3 smooth; scutellar apical projections dorsally DIAGNOSIS: Males recognized among other directed (pl. 15C); meso- and metasterna sepaspecies in Gibbosella by the circular head shape rated by distinct transverse suture (pl. 15F); in lateral view that is as long as it is wide in dor- mesosternum with large medial depression; sal view, synthlipsis less than the width of an eye, metasternum slightly medially longitudinally large ocelli, eyes reaching dorsal and ventral depressed; forefemur incrassate; fossula spongihead margin, labial segment II longer than III, osa on mid tibia; tarsomeres I and II combined mesosternum with large medial depression, fos- shorter than III; hemelytron surpassing abdomisula spongiosa on the mid tibia, and BPE as long nal apex; distal part of R present but not forming as the basal plate. This species is similar to G. R+M cell (pl. 17B, C). ABDOMEN: Apex medially betampona , but the slightly darker pronotum, notched; sternites medially convex, intersegmenpale meso- and metasterna and corium, smaller tal sutures carinulate between II–IV and laterally synthlipsis width, meso- and metasterna sepa- between IV–VI; pygophore process subtriangular rated by a distinct suture, presence of the fossula in lateral (pl. 20A) and caudal (pl. 19G, I) views, spongiosa on the mid tibia, and longer BPE rela- apex rounded; BPE as long as basal plate; endotive to the basal plate distinguish this species somal struts reaching posterior margin of DPS, from G. betampona . with posterior ventral process (pl. 22G); area of DESCRIPTION: MALE: Body length: 5.42 mm endosomal struts–DPS fusion basally narrow but (holotype). COLORATION: Dark brown with distally wider and ovate (pl. 21A). pale labial segment IV, apical half of antennal ETYMOLOGY: The species epithet is a noun in segment V, antennal segments VI–VIII, posterior apposition and is named after the type locality, margin of pronotum, stridulatory groove, meso- Camp Mantella, Marojejy National Park, and metasterna, corium and basal wing veins, Madagascar. coxae, trochanters, femora basally, mid and hind DISTRIBUTION: Marojejy Nature Reserve in tibiae medially, tarsi, laterotergites except poste- Antsiranana province (map 3). rior margin, and sternites medially and laterally. HOLOTYPE: Male: Madagascar: Antsiranana: Remainder of antenna and labium, dorsal surface Marojejy National Park, 5 km W Manantenina of pronotum, and legs brown. VESTITURE: village, Camp Mantella, 14.43816 ° S 49.774 ° E, Sparse; other features as in generic description. 490 m, 11 Feb 2005 – 18 Feb 2005, M. Irwin, R. STRUCTURE: HEAD (pl. 13C): Ovoid in lateral Harin’Hala (00006458) (CAS).

46 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

PARATYPES: Madagascar: Antsiranana: Marojejy antenna 8-segmented ; labium stout; labial segment National Park, 5 km W Manantenina village, Camp III subequal to II, ventrally convex (pl. 13C). THO- Mantella , 14.43816 ° S 49.774 ° E, 490 m, 28 Apr 2005 –07 RAX: Anterior pronotal lobe with very small antero-

May 2005, M. Irwin, R. Harin’Hala, 13 (00006460) ( CAS) ; 14 Oct 2005 – 22 Oct 2005, M. Irwin, R. lateral projections (pl. 15B) ; pronotal transverse Harin’Hala, 13 (00007038) ( UCR). R. N.I. de Marojejy, furrow continuous ; lateral depressions on posterior 10.0 km NW Manantenina , 14.43333 ° S 49.76167 ° E, 750 pronotal lobe smooth ; scutellar apical projections m, 15 Oct 1996 – 22 Oct 1996, E. Quinter and T. Nguyen , dorsally directed (pl. 15C) ; separation of meso- and 13 (00078367) (AMNH). metasterna obscured; mesosternum with medial and paramedial longitudinal depressions; metaster-

Gibbosella megafrons , new species num slightly medially longitudinally depressed;

Plates 1, 5, 8; map 1 forefemur slightly incrassate; presence of fossula

DIAGNOSIS: Males recognized among other spongiosa on mid tibia cannot be determined due species in Gibbosella by the very small body size, to tibial apex obscured; tarsomeres I and II comcoloration, anteocular region shorter than the bined shorter than III; hemelytron surpassing postocular, postclypeus depressed, synthlipsis abdominal apex; distal part of R absent. ABDOMEN: width about 3.5 times the width of an eye, and Apex medially notched; sternites medially convex, distal part of R absent. This species is similar to intersegmental sutures carinulate between II–VI; G. andasibe , from which it differs by the entirely pygophore process subtriangular in lateral (pl. 20A) dark brown pronotum and scutellum, depressed and caudal (pl. 19G, I) views, apex rounded; basal postclypeus, larger synthlipsis width, presence of plate and BPE damaged; endosomal struts reaching a fossula spongiosa on the mid tibia, and inter- posterior margin of DPS, with posterior ventral segmental sutures carinulate between II–VI. process (pl. 22G); area of endosomal struts–DPS

DESCRIPTION: MALE: Body length: 3.78 mm fusion elongate subquadrate. (holotype), 3.78–5.05 mm. COLORATION: Dark ETYMOLOGY: The species epithet is a noun in brown with pale antennal segments VI and VII, the nominative case and is named for the very posterodorsal spot on mesopleuron, spots on large synthlipsis width. pleura dorsad of coxae, corium basally, coxae, tro- DISTRIBUTION: Ranomafana National Park chanters, femora basally, anterior half of lateroter- and Midongy du Sud National Park in the Fiangites , sternites III – VI medially, and anterolateral arantsoa province (map 1). margins of sternites pale. Antennal segments IV HOLOTYPE: Male : Madagascar: Fianarant- and V, labium, scutellar processes, remainder of soa: Parc National Ranomafana, Vohiparara, at corium, basal wing veins, and remainder of legs broken bridge, 21.22616 ° S 47.36983 ° E, 1110 m, brown. Wing membrane light brown. VESTI- 28 Nov 2001 – 06 Dec 2001, M. Irwin, R.

TURE: Sparse ; other features as in generic descrip- Harin’Hala (00007168) ( CAS). tion. STRUCTURE: HEAD: Ovoid in lateral view PARATYPE: Madagascar: Fianarantsoa: Sud-Est (pl. 13F) ; longer than wide in dorsal view; anteocu- Midongy, Sud Mont Papango (sud-ouest Befotaka ), 23.85000 ° S

46.98333

° E, 1200 m, Mar 1959, Andria

lar region shorter than postocular; gula flat, con- Robinson, 13 (MNHN).

forming to rounded shape of head (pl. 13C); postclypeus with shallow, broad medial longitudi- Gibbosella mirabilis Chłond, 2010 nal depression to middle of interocular area (pl. Plate 12; map 1 13B); synthlipsis about 3.5 times width of eye; Gibbosella mirabilis Chłond, 2010, 2522: 63 .

interocular sulcus posterior to hind margin of eye; ocelli small, located on shallow median tubercle (pl. DISTRIBUTION: The only known specimen 13I); eye about one-fourth of head length, not was collected from “Périnet” (currently known as reaching dorsal and ventral head surfaces; antenna Andasibe-Mantadia National Park) in the inserted dorsally on head; pedicel as long as scape; Toamasina province (map 1).

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 47

DISCUSSION: This species was recently view; anteocular region as long as postocular (pl. described by Chłond (2010a) from a single 13C); gula slightly swollen ventrolaterally, not female representative. No other female or male distinctly produced beyond ventral head margin specimens have been examined, and, thus, this (pl. 13F–K); postclypeus with deep, narrow species is not redescribed. This species is very medial longitudinal depression to middle of similar to G. elongata , but is distinguished by the interocular area; synthlipsis two times width of more strongly dorsally elevated anterior pronotal eye; interocular sulcus near hind margin of eye lobe, convergent scutellar processes, and hind (pl. 13B); ocelli large, located on shallow median tibiae with dark coloration on the apical half. tubercle (pl. 13I); eye about one-fourth of head HOLOTYPE: Female : Madagascar: Toama- length, not reaching dorsal and ventral head sursina: Périnet (Andasibe), 18.82666 ° S 48.44778 ° E, faces; antenna inserted sublaterally on head; 1119 m, Oct 1972, A. Peyrieras ( MNHN). pedicel slightly longer than scape; antenna 8-seg-

mented; labium stout; labial segment III shorter

Gibbosella nitida , new species than II, ventrally convex (pl. 13C). THORAX:

Plates 1, 5; map 2 Anterior pronotal lobe without anterolateral pro- DIAGNOSIS: Males are recognized among jections; pronotal transverse furrow continuous; other species in this genus by the small body lateral depressions on posterior pronotal lobe size, synthlipsis two times the width of an eye, smooth; scutellar apical projections dorsally eye one-fourth of head length, fossula spongiosa directed (pl. 15C); ventrally obscured by card absent on the mid tibia, and corium with rela- mount; forefemur incrassate; fossula spongiosa tively large shiny granulations not covered by absent on mid tibia; tarsomeres I and II comsetae, as well as small granulations with very bined shorter than III; hemelytron surpassing short, erected, flattened setae. This species is abdominal apex. ABDOMEN: Apex rounded; stersimilar to G. megafrons , but is distinguished by nites medially obscured by card mount, but the smaller body size, more subpentagonal head intersegmental sutures carinulate at least laterally that is as long as it is wide, anteocular length between II–VI; pygophore process subtriangular subequal to postocular, smaller synthlipsis width, in lateral (pl. 20A) and caudal (pl. 19G, I) views, longer labial segment II relative to III, and apex rounded; BPE as long as basal plate; area of appearance of the corium. endosomal struts–DPS fusion basally narrow but DESCRIPTION: MALE: Body length: 4.75 mm distally wider and ovate subquadrate. (holotype). COLORATION: Dark brown with ETYMOLOGY: The species epithet is an adjeclabial segment III and IV brown. Labial segment tive in the nominative case and is named for the II apically and III basally, basal two-thirds of shiny hemelytra. scape, coxae, trochanters, femora ventrally, and DISTRIBUTION: The only known specimen sternites II–VI medially and laterally pale. was collected in the Marojejy Nature Reserve in Corium with shiny areas due to relatively large that Antsiranana province (map 2). shiny granulations not covered by setae, as well HOLOTYPE: Male: Madagascar: Antsiranana: as dull areas due to very small granulations with Sambava District, Marojejy, Ambinanitelo, very short, erect, flattened setae. VESTITURE: 14.44133 ° S 49.73908 ° E, 500 m, Dec 1958, Raha- Dense; corium with shorter, flattened setae; other rizonina (MNHN). features as in generic description. Hemelytra covered by very dense, small granulations with Gibbosella notoconica , new species very short and flattened setae, as well as sparse Plates 1, 5, 8, 20A, 21A; map 2 relatively large granulations without setae. DIAGNOSIS: Males are recognized among other STRUCTURE: HEAD: Ovate, almost pentagonal species in this genus by a combination of the folin lateral view; about as long as wide in dorsal lowing characters: the larger body size, synthlipsis

48 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

as wide as an eye, large ocelli, eyes reaching the antenna inserted sublaterally on head; pedicel dorsal and ventral head margins in lateral view, about one-fourth longer than scape; antenna labial segment III ventrally straight, conically ele- 8-segmented; labium slender; labial segment III vated anterior pronotal paramedian lobes in lat- subequal to II, ventrally straight (pl. 13G–I, K). eral view, anterior pronotal lobe without THORAX: Anterior pronotal lobe conical in latanterolateral protuberances, continuous pronotal eral view (pl. 15C), without distinct anterolateral transverse suture, and mesosternum with large projections; pronotal transverse furrow continumedial depression. This species is similar to G. ous; lateral depressions on posterior pronotal conisimilis , but the slightly larger body size, pale lobe transversely striated; scutellar apical projecantennal segment V, dark brown pronotum with tions dorsally directed (pl. 15C); meso- and pale lateral spots extending from posterior half of metasterna separated by distinct transverse anterior lobe to anterior half of posterior lobe, suture (pl. 15F); mesosternum with large medial pale meso- and metasterna, smaller synthlipsis depression; metasternum slightly medially longiwidth, interocular sulcus near the hind margin of tudinally depressed; legs slender; fossula sponthe eye, larger ocelli, eye size relative to the head giosa on mid tibia; tarsomeres I and II combined length and height, the more distinctive conical subequal to III; hemelytron surpassing abdomianterior pronotal paramedian lobes with a dorsal nal apex; distal part of R present but not forming protuberance, and intersegmental sutures carinu- R+M cell (pl. 17B, C). ABDOMEN: Apex medially late between II–IV and laterally between IV–VI notched or sinuate; sternites medially convex, distinguish this species from G. conisimilis . intersegmental sutures carinulate between II–IV DESCRIPTION: MALE: Body length: 9.48 mm and laterally between IV–VI; pygophore process (holotype), 8.53–9.48 mm. COLORATION: subtriangular in lateral (pl. 20A) and caudal (pl. Dark brown with pale synthlipsis laterally, post- 19G, I) views, apex rounded; BPE as long as ocular dorsally except ocellar tubercle, neck dor- basal plate; endosomal struts reaching posterior sally, scape basally, antennal segments V–VIII, margin of DPS, with posterior ventral process anterior and posterolateral margins of pronotum, (pl. 22G); area of endosomal struts–DPS fusion lateral spots extending from posterior half of ovate (pl. 21A). anterior pronotal lobe to anterior half of poste- ETYMOLOGY: The species epithet is an adjecrior lobe, stridulatory groove, meso- and tive in the nominative case and is named for the metasterna, coxae, trochanters, forefemur basally conical anterior pronotal paramedian lobes. and ventrally, mid and hind femora basally and DISTRIBUTION: Marojejy Nature Reserve in subapically, tibiae medially, anterior area of lat- Antsiranana province (map 2). erotergites, lateral margins of sternites, and ster- DISCUSSION: The coloration is slightly variable nites medially. Labium and remainder of antenna with the postclypeus entirely dark brown, the brown. VESTITURE: Sparse; other features as in pronotum less pale, and the brown ventrolateral generic description. STRUCTURE: HEAD: Ovoid longitudinal band on the sternites varying in size in lateral view (pl. 13F); longer than wide in dor- in some specimens. sal view; anteocular region as long as postocular HOLOTYPE: Male: Madagascar: Antsiranana: (pl. 13C); gula flat, conforming to rounded shape Marojejy National Park, 5 km W Manantenina of head (pl. 13C); postclypeus with deep, narrow village, Camp Mantella, 14.43816 ° S 49.774 ° E, medial longitudinal depression to middle of 490 m, 30 May 2005 – 11 Jun 2005, M. Irwin, R. interocular area; synthlipsis about width of eye; Harin’Hala (00045367) (CAS). interocular sulcus near hind margin of eye (pl. PARATYPES: Madagascar: Antsiranana: Marojejy 13B); ocelli large, located on distinct median National Park, 5 km W Manantenina village, Camp tubercle (pl. 13C); eye about half of head length, Mantella, 14.43816 ° S 49.774 ° E, 490 m, 25 Dec 2004 –30 reaching dorsal and ventral head surfaces; Dec 2004, M. Irwin, R. Harin’Hala, 13 (00007240)

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 49

(CAS); 18 Feb 2005 – 25 Feb 2005, M. Irwin, R. description. STRUCTURE: HEAD: Circular in lat- Harin’Hala, 13 (00045655) (CAS); 11 Mar 2005 –18 eral view (pl. 13C); as long as wide in dorsal view;

Mar 2005, M. Irwin, R. Harin’Hala,

13

(00044813)

anteocular region as long as postocular (pl. 13C); (SU); 28 Apr 2005 – 07 May 2005, M. Irwin, R .

Harin’Hala, 13 (00007174) (UCR); 14 Oct 2005 –22 gula flat, conforming to rounded shape of head (pl. Oct 2005, M. Irwin, R. Harin’Hala, 13 (00045558) 13C); postclypeus with deep, narrow medial longi- (UCR). Unknown: 13 (00006144) (CAS). tudinal depression to middle of interocular area;

synthlipsis about width of eye; interocular sulcus Gibbosella pallidacorium , new species near hind margin of eye (pl. 13B); ocelli large,

Plates 1, 5, 8; map 1 located on shallow median tubercle (pl. 13I); eye DIAGNOSIS: Males are recognized among about half of head length, nearly reaching dorsal other species of Gibbosella by a combination of and ventral head surface; antenna inserted dorsally the following characters: the coloration, circular on head; pedicel about one-fourth longer than head shape in lateral view that is as long as it is scape; antenna 8-segmented; labium slender; labial wide in dorsal view, large ocelli, anterior prono- segment III subequal to II, ventrally convex (pl. tal lobe laterally carinate, meso- and metasterna 13C). THORAX: Anterior pronotal lobe with very not completely separated by a distinct suture, small anterolateral projections (pl. 15B), slightly and intersegmental sutures carinulate between carinate laterally; pronotal transverse furrow con- II–IV and laterally between IV and V. This spe- tinuous; lateral depressions on posterior pronotal cies is similar to G. betampona , but differs by the lobe transversely striated; scutellar apical projecpale postocular and neck, pale markings on the tions dorsally directed (pl. 15C); meso- and pronotum, pale corium, pale ventral abdominal metasterna not completely separated by distinct surface with dark brown longitudinal stripe lat- transverse suture (pl. 15F); mesosternum with erally, circular head shape, relatively deeper medial and paramedial longitudinal depressions; postclypeal depression, eye not reaching ventral metasternum slightly medially longitudinally head margin, labial segment II and III subequal, depressed; forelegs missing, mid and hind legs anterior pronotal lobe laterally carinate, meso- slender; fossula spongiosa on mid tibia; tarsi misssternum with three longitudinal depressions, fos- ing; hemelytron surpassing abdominal apex; distal sula spongiosa on the mid tibia, and part of R absent. ABDOMEN: Apex sinuate; sternites intersegmental sutures carinulate between II–IV medially convex, intersegmental sutures carinulate and laterally between IV and V. between II–IV and laterally between IV and V; DESCRIPTION: MALE: Body length: 5.78 mm pygophore process subtriangular in lateral (pl. (holotype). COLORATION: Dark brown with 20A) and caudal (pl. 19G, I) views, apex rounded; pale postocular and neck dorsally, scape basally, BPE as long as basal plate; endosomal struts reachapical half of antennal segment V and segments VI ing posterior margin of DPS, posteriorly obscured and VII, labial segment IV, anterior pronotal lobe by semiextended endosoma; area of endosomal anteromedially, pronotal disc, posterior pronotal struts–DPS fusion circular.

lobe anterolaterally and posterolateral margin, ETYMOLOGY: The species epithet is a noun in stridulatory groove, meso- and metasterna, corium the nominative case and is named for the pale except apical margin, basal wing veins, coxae, tro- corium.

chanters, mid and hind femora basally and medi- DISTRIBUTION: The only known specimen ally (forelegs missing), mid tibia medially and was collected in Marojejy Nature Reserve in apically, hind tibia except basally, dorsal lateroter- Antsiranana province (map 1).

gites except posterior margin, ventral lateroter- HOLOTYPE: Male: Madagascar: Antsiranana: gites, and sternites except pygophore and laterally. R. N.I. de Marojejy, 11.0 km NW Manantenina, Labial segment III and wing membrane brown. 14.43667 ° S 49.74167 ° E, 1225 m, 25 Oct 1996 –03 VESTITURE: Sparse; other features as in generic Nov 1996, Eric L. Quinter (00078364) ( AMNH).

50 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

Gibbosella pallidalata , new species paramedial longitudinal depressions; metaster- Plates 1, 5, 8; map 1 num slightly medially longitudinally depressed; DIAGNOSIS: Males recognized among other forefemur slightly incrassate ; fossula spongiosa species in Gibbosella by the coloration, 6-seg- absent on mid tibia; tarsomeres I and II commented antenna, anterior pronotal lobe without bined subequal to III on forelegs, longer on mid distinct anterolateral protuberances, and fossula and hind legs; hemelytron surpassing abdominal spongiosa absent on the mid tibia. This species is apex; distal part of R present but not forming similar to G. planiscutum , but, in addition to the R +M cell (pl. 17B, C). ABDOMEN: Apex rounded diagnostic features, it differs by the continuous or sinuate ; sternites medially convex, intersegpronotal transverse suture, separation of the mental sutures carinulate between II–VI; pygomeso- and metasterna, and intersegmental phore process flattened in lateral view (pl. 20F), sutures carinulate between II – VI. subtriangular in caudal view (pl. 19G, I), apex DESCRIPTION: MALE: Body length: 5.23 mm rounded ; BPE shorter than basal plate; endo- (holotype), 4.88–5.34 mm. COLORATION: Pale somal struts not reaching posterior margin of with dark brown head, propleuron medially, DPS, posteriorly obscured by semiextended mesopleuron and metapleuron except dorsad of endosoma; area of endosomal struts–DPS fusion coxae, thoracic sterna, sternites ventrolaterally, elongate subquadrate. and pygophore medially. Antenna, labium, pos- ETYMOLOGY: The species epithet is a noun in terior pronotal lobe medially, scutellum, distal the nominative case and is named for the pale half of corium, legs except femora basally and wing membrane. ventrally and mid and hind tibiae medially, and DISTRIBUTION: Fianarantsoa and Toliara posterior half of laterotergites II – VI brown. provinces (map 1). VESTITURE: Sparse ; other features as in generic DISCUSSION: Some specimens have small description. STRUCTURE: HEAD: Ovoid in lat- paramedian brown spots on the anterior pronoeral view (pl. 13F) ; longer than wide in dorsal tal lobe. view; anteocular region as long as postocular (pl. HOLOTYPE: Male : Madagascar: Fianarant- 13C) ; gula moderately swollen ventrolaterally, soa: Parc National Ranomafana, Belle Vue at not distinctly produced beyond ventral head Talatakely, 21.2665 ° S 47.42016 ° E, 1020 m, 14 Apr margin (pl. 13F–K) ; postclypeus with shallow, 2002– 23 Apr 2002, M. Irwin, R. Harin’Hala narrow medial longitudinal depression to middle (00044983) ( CAS). of interocular area ; synthlipsis about two times PARATYPES: Madagascar: Fianarantsoa: Parc width of eye ; interocular sulcus posterior to hind National Ranomafana, Belle Vue at Talatakely, 21.2665 ° S 47.42016 ° E, 1020 m, 16 Oct 2001 –08 Nov margin of eye ; ocelli small, located on shallow 2001, M. Irwin, R. Harin’Hala, 23 (00007237, median tubercle (pl. 13I); eye about one-fourth 00007245) ( CAS) ; 08 Nov 2001 – 15 Nov 2001, M. of head length, not reaching dorsal and ventral Irwin, R. Harin’Hala , 13 (00007159) ( SU) ; 22 Nov head surfaces; antenna inserted sublaterally on 2001– 28 Nov 2001, M. Irwin, R. Harin’Hala , 23 head ; pedicel about as long as scape; antenna (00045117, 00045624) (BMNH); 04 Feb 2002 – 12 Feb 2002, M. Irwin, R. Harin’Hala , 13 (00007206) ( CAS) ; 6-segmented; labium stout; labial segment III 26 Feb 2002 – 04 Mar 2002, M. Irwin, R. Harin’Hala, shorter than II, ventrally convex (pl. 13C). THO- 13 (00007130) ( AMNH) ; 31 Mar 2002 – 07 Apr 2002, RAX: Anterior pronotal lobe without distinct M. Irwin, R. Harin’Hala , 23 (00045535, 00045541) anterolateral projections ; pronotal transverse (USNM); 07 Apr 2002 – 14 Apr 2002, M. Irwin, R. furrow continuous ; lateral depressions on poste- Harin’Hala, 13 (00044918) ( CAS) ; 14 Apr 2002 –23 rior pronotal lobe smooth; scutellar apical pro- Apr 2002, M. Irwin, R. Harin’Hala, 23 (00007074, 00044825) ( MNHN), 23 (00044914, 00045473) jections dorsally directed (pl. 15C) ; meso- and (BMNH); 28 Apr 2002 – 05 May 2002, M. Irwin, R. metasterna separated by distinct transverse Harin’Hala , 23 (00005356, 00007215) ( SU) ; 05 May suture (pl. 15F); mesosternum with medial and 2002– 13 May 2002, M. Irwin, R. Harin’Hala , 13

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 51

(00007232) (CAS); 04 Jul 2002 – 14 Jul 2002, M. Irwin, the following characters: the coloration, dense

R. Harin’Hala, 33 (00007205, 00007222, 00007243) vestiture, pronotal transverse suture divided by

(CAS); 14 Jul 2002 – 24 Jul 2002, M. Irwin, R. paramedian ridges, scutellar processes horizon-

Harin’Hala, 13 (00007246) (UCR). Parc National

Ranomafana, Vohiparara, at broken bridge, tally directed, meso- and metasterna not entirely

21.22616 ° S 47.36983 ° E, 1110 m, 28 Nov 2001 –06 Dec separated by a distinct suture, and intersegmen-

2001, M. Irwin, R. Harin’Hala, 13 (00044934) (CAS), tal sutures carinulate between II–VI and laterally

13 (00045471) (MNHN); 15 Dec 2001 – 21 Dec 2001, between VI and VII. Females are recognized by

M. Irwin, R. Harin’Hala, 13 (00099046) (CAS); 28 the coloration and synthlipsis about three times

Jan 2002– 04 Feb 2002, M. Irwin, R. Harin’Hala, 13 the width of an eye. Males are similar to G.

(00044827) (SU); 04 Feb 2002 – 12 Feb 2002, M. Irwin,

R. Harin’Hala, 13 (00007235) (CAS); 06 May 2002 – brunalvus , from which it differs by the pale

15 May 2002, M. Irwin, R. Harin’Hala, 13 (00007194) markings on the head (e.g., mandibular and

(CAS); 04 Jun 2002 – 14 Jun 2002, M. Irwin, R. maxillary plates, antennifers, ventral and lateral

Harin’Hala, 23 (00007262, 00007267) (USNM); 15 anteocular and postocular surface), pale prono-

Jul 2002– 25 Jul 2002, M. Irwin, R. Harin’Hala, 13 tum with dark brown paramedian spots on the

(00045533) (UCR). Parc National Ranomafana, radio

tower at forest edge, 21.251 ° S 47.40716 ° E, 1130 m, 28 anterior lobe and margins of posterior lobe,

Nov 2001– 06 Dec 2001, M. Irwin, R. Harin’Hala, 13 depressed postclypeus, orientation of the scutel-

(00045580) (CAS); 31 Mar 2002 – 09 Apr 2002, M. lar processes, lack of a distinct separation

Irwin, R. Harin’Hala, 13 (00007132) (CAS); 09 Apr between the meso- and metasterna, and interseg-

2002– 16 Apr 2002, M. Irwin, R. Harin’Hala, 13 mental sutures carinulate between II–VI and lat-

(00007228) (AMNH); 15 Jul 2002 – 25 Jul 2002, M. erally between VI and VII. Females are very

Irwin, R. Harin’Hala, 13 (00005358) (USNM); 07

Sep 2003– 18 Sep 2003, M. Irwin, R. Harin’Hala, 23 similar to G. brunalvus , but are distinguished by

(00007108, 00044902) (CAS); 21 Mar 2004 –02 Apr the nearly black body, pale apex of antennal seg-

2004, M. Irwin, R. Harin’Hala, 13 (00007036) (SU); ment VIII and apex of scutellum, and larger

18 Mar 2006 – 30 Mar 2006, M. Irwin, R. Harin’Hala, synthlipsis width.

13 (00007145) (CAS); 17 Oct 2006 – 01 Nov 2006, M. DESCRIPTION: MALE: Body length: 5.08 mm

Irwin, R. Harin’Hala, 13 (00045653) (UCR). Rano- (holotype), 4.91–5.90 mm. COLORATION:

mafana JIRAMA water works, 21.2485 ° S 47.45216 ° E,

690 m, 22 Nov 2001 – 28 Nov 2001, M. Irwin, R. Dark brown with pale mandibular and maxillary

Harin’Hala, 13 (00007210) (CAS), 13 (00007227) plates, antennifers, ventral and lateral anteocular

(UCR). Ranomafana National Park, Talatakely, and postocular surface, except at posterior mar-

21.25 ° S 47.41667 ° E, 900 m, 09 Jan 2001 – 19 Jan 2001, gin, scape basally, antennal segments VI–VIII,

D.H. and K.M. Kavanaugh, R.L. Brett, E. Elsom, and anterior pronotal lobe except paramedian spots

F. Vargas, 23 (00007110, 00044853) (CAS), 23

(00044979, 00044996) (AMNH). Toliara: Berenty and pronotal longitudinal furrow, posterior pro-

Special Reserve, 8 km NW Amboasary, 25.021 ° S notal lobe except medially, anterior propleural

46.3055 ° E, 35 m, 29 Aug 2004 – 08 Sep 2004, M. Irwin, lobe except medially, posterior propleural lobe

F. Parker, R. Harin’Hala, 13 (00044876) (CAS). Parc dorsally, mesopleuron and metapleuron dorsad

National d’Andohahela, Col du Sedro, 3.8 km 113° of coxae, apical half of scutellum, basal half of

ESE Mahamavo, 37.6 km 341° NNW Tolagnaro, corium, coxae, trochanters, forefemur basally

24.76388 ° S 46.75166 ° E, 900 m, 21 Jan 2002 –25 Jan

2002, Fisher et al., 13 (00006463) (UCR), 13 and ventrally, mid and hind femora basally, tib-

(00007085) (CAS), 13 (00007247) (MNHN). iae basally, anterior half of laterotergites, and

Unknown: 63 (00007076, 00007125, 00007204, sternites medially and anterolaterally. Remainder

00007216, 00007249, 00007264) (CAS). of antenna, labium, remainder of legs, apical half of corium, and basal wing veins brown. VESTI-

Gibbosella planiscutum , new species TURE: Dense; other features as in generic Plates 1, 5, 8, 17A, 23B; map 3 description. STRUCTURE: HEAD: Ovoid in lat-

DIAGNOSIS: Males are recognized among eral view (pl. 13F); longer than wide in dorsal

other species in this genus by a combination of view; anteocular region as long as postocular (pl.

52 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

13C); gula moderately swollen ventrolaterally, brown. STRUCTURE: In addition to characters not distinctly produced beyond ventral head mentioned in the generic description: HEAD: margin (pl. 13F–K) ; postclypeus with shallow, Synthlipsis about three times width of eye. THObroad medial longitudinal depression to middle RAX: Pronotal longitudinal furrow restricted to of interocular area (pl. 13B) ; synthlipsis about anterior lobe; lateral depressions on posterior two times width of eye; interocular sulcus poste- pronotal lobe obsolete; forefemur with small rior to hind margin of eye; ocelli small, located ventral tubercles (pl. 16D). ABDOMEN: Intersegon shallow median tubercle (pl. 13I) ; eye about mental sutures carinulate between sternites II– one-fourth of head length, not reaching dorsal VI; external genitalia as in plate 23B; bursa and ventral head surfaces; antenna inserted dor- copulatrix membranous, with lateral lobes (pl. sally on head; pedicel about as long as scape; 23F); vermiform gland elongate, much longer antenna 8-segmented; labium stout; labial seg- than bursa copulatrix; lateral spermathecae ment III shorter than II, ventrally convex (pl. inserted on median oviduct with slender, fragile 13C). THORAX: Anterior pronotal lobe with very duct and large ovoid pouch. small anterolateral projections (pl. 15B) ; pronotal ETYMOLOGY: The species epithet is a noun in transverse furrow divided by paramedian longi- the nominative case and is named for the horitudinal ridges (pl. 15B); lateral depressions on zontally oriented scutellar apices. posterior pronotal lobe transversely striated; scu- DISTRIBUTION: Antananarivo and Mahajanga tellar apical projections horizontally directed (pl. provinces (map 3). 15D) ; meso- and metasterna not completely DISCUSSION: In some specimens, the entire separated by distinct transverse suture (pl. 15E) ; postocular is dark brown, the brown spots on the mesosternum with medial and paramedial longi- paramedian anterior pronotal lobes are absent, tudinal depressions; metasternum slightly medi- and the scutellum is entirely pale. Females were ally longitudinally depressed; fore- and mid associated to males based on morphology and femora incrassate; fossula spongiosa on mid the same collection event as another male. tibia; tarsomeres I and II combined shorter than HOLOTYPE: Male : Madagascar: Mahajanga: III ; hemelytron reaching or surpassing abdomi- Analamanitra Forest , 14 km NE of Misinjo, nal apex ; distal part of R vein present but not 16.13333 ° S 45.7 ° E, 20 m, 20 Nov 2007 –04 Dec forming R +M cell (pl. 17B, C). ABDOMEN: Apex 2007, M. Irwin, R. Harin’Hala (00044849) ( CAS). rounded ; sternites with shallow medial depres- PARATYPES: Madagascar: Antananarivo: 46 km NE sion (pl. 18C), intersegmental sutures carinulate of Ankazobe: Ambohitantely , 18.198 ° S 47.2815 ° E, 700 m, 10 Jan 2004 – 20 Jan 2004, M. Irwin, R. Harin’Hala, 13 between II – VI and laterally between VI and VII; (00006461) ( CAS) , 13 (00007266) (SU); 20 Jan 2004 –01 pygophore process subtriangular in lateral (pl. Feb 2004, M. Irwin, R. Harin’Hala, 13 (00007058) 20A) and caudal (pl. 19G, I) views, apex rounded; ( UCR) . Mahajanga: 160 km N of Maevatanana on RN BPE shorter than basal plate ; endosomal struts 04, Ampijoroa National Park, 16.31933 ° S 46.81333 ° E, 43 reaching posterior margin of DPS, with posterior m, 07 Feb 2005 – 19 Feb 2005, M. Irwin, R. Harin’Hala, 13 (00045216) ( USNM). Namoroka, 53 km from Soaventral process (pl. 22G), fused anterior to fusion lala, 3 km N Vilanandro Village, 16.47333 ° S 45.39133 ° E, with DPS ; area of endosomal struts–DPS fusion 122 m, 30 Nov 2007 – 09 Dec 2007, M. Irwin, R. ovate (pl. 21A). FEMALE: Body length: 4.78– Harin’Hala, 13 (00045296) ( USNM). Parc National de 5.06 mm. Similar to males, but differ in the fol- Namoroka , 16.9 km 317° NW Vilanandro, 16.40666 ° S lowing characters: COLORATION: Dark brown 45.31 ° E, 100 m, 12 Nov 2002 – 16 Nov 2002, Fisher et al., with brown scape, pedicel, antennal segments 13 (00006320), 1♀ (00045373) ( CAS), 1♀ (00045711) ( UCR). Parc National de Namoroka , 17.8 km 329° III – IV, base of segment V, apex of segment VIII, WNW Vilanandro , 16.37666 ° S 45.32666 ° E, 100 m, 08 labium, and legs except trochanter and tarsi. Nov 2002– 12 Nov 2002, Fisher et al., 13 (00007209) Remainder of antennal segments, scutellar pro- ( CAS). Réserve forestière Beanka , 50.2 km E Maintirano, cesses, trochanters, and tarsi whitish to pale 18.02638 ° S 44.05055 ° E, 250 m, 19 Oct 2009 –26 Oct

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 53

2009, B.L. Fisher et al., 13 (00006464) ( CAS). Réserve margin of eye ; ocelli small, located on shallow Spéciale de Bemarivo , 23.8 km 223° SW Besalampy, median tubercle (pl. 13I) ; eye about one-fourth 16.925 ° S 44.36833 ° E, 30 m, 19 Nov 2002 – 23 Nov 2002, of head length, not reaching dorsal and ventral Fisher et al., 13 (00045515) (SU). Unknown : 13

head surfaces; antenna inserted dorsally on head; (00007149) (CAS).

pedicel about as long as scape; antenna 8-seg-

mented; labium stout; labial segment III sub- Gibbosella quadocris , new species equal to II, ventrally convex (pl. 13C). THORAX: Plates 1, 5, 8, 19A, 20B; map 2 Anterior pronotal lobe with very small anterolat- DIAGNOSIS: Males are recognized among eral projections (pl. 15B); pronotal transverse other species in Gibbosella by the small body furrow continuous; lateral depressions on postesize, ovoid head shape in lateral view that is lon- rior pronotal lobe smooth; scutellar apical proger than it is wide in dorsal view, synthlipsis 1.5 jections dorsally directed (pl. 15C); meso- and times the width of an eye, labial segments II and metasterna not completely or weakly separated III subequal, subquadrate median pygophore by distinct transverse suture (pl. 15E); mesosterprocess with truncated apex in caudal view, and num with medial and paramedial depressions; BPE as long as the basal plate. This species is metasternum slightly medially longitudinally very similar to G. vangocris , but is distinguished depressed; forefemur incrassate; fossula spongiby the smaller body size, slightly shorter head, osa on mid tibia; tarsomeres I and II combined more narrowly depressed postclypeus, relative subequal to III; hemelytron surpassing abdomilengths of the labial segments, meso- and nal apex; distal part of R present but not forming metasterna not distinctly or weakly separated by R+M cell (pl. 17B, C). ABDOMEN: Apex rounded a distinct transverse suture, and shape of the or sinuate; sternites longitudinally depressed pygophore process in caudal view. medially (pl. 18C), intersegmental sutures cari- DESCRIPTION: MALE: Body length: 5.39 mm nulate between II–VI; pygophore (pls. 19A, 20B) (holotype), 5.33–5.78 mm. COLORATION: process subtriangular in lateral view, subquad- Dark brown with pale interocular sulcus, ante- rate in caudal view, apex truncated; BPE as long rior pronotal lobe except interspersed brown as basal plate; endosomal struts not reaching markings posterolateral margin of pronotum, posterior margin, posteriorly obscured by metasternum, corium basally, coxae, trochanters, semiextended endosoma; fused anterior to fusion femora basally and ventrally, mid and hind tibiae with DPS; area of endosomal struts–DPS fusion basally, laterotergites except posterior margin, elongate ovoid.

and sternites (except medial pygophore and lat- ETYMOLOGY: The species epithet is a noun in eral and posterolateral margins of sternites II– apposition and is named for the subquadrate VI). Antennal segment V (except basally) and median pygophore process.

VI–VIII white. Remainder of antenna, labium, DISTRIBUTION: Ranomafana National Park, parts of anterior pronotal lobe, posterior prono- Fianarantsoa province (map 2).

tal lobe, and rest of corium and basal wing veins DISCUSSION: In some specimens, the postebrown. VESTITURE: Sparse ; other features as in rior pronotal lobe is dark brown, the anterior generic description. STRUCTURE: HEAD: Ovoid pronotal lobe has reduced pale coloration and in lateral view (pl. 13F); longer than wide in dor- appears almost uniformly brown, and the corium sal view; anteocular region as long as postocular and basal wing veins range from pale to brown. (pl. 13C); gula flat, conforming to rounded shape HOLOTYPE: Male : Madagascar: Fianarantof head (pl. 13C); postclypeus with shallow, nar- soa: Parc National Ranomafana , Belle Vue at row medial longitudinal depression to middle of Talatakely, 21.2665 ° S 47.42016 ° E, 1020 m, 08 interocular area; synthlipsis about 1.5 times Nov 2001– 15 Nov 2001, M. Irwin, R. Harin’Hala width of eye; interocular sulcus posterior to hind (00007072) ( CAS).

54 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

PARATYPES: Madagascar: Fianarantsoa: Parc segment III shorter than II, intersegmental National Ranomafana, Belle Vue at Talatakely, 21.2665 ° S sutures carinulate between II–VI, and spadelike 47.42016 ° E, 1020 m, 08 Nov 2001 – 15 Nov 2001, M. median pygophore process in caudal view. This

Irwin, R. Harin’Hala, 23 (00006457, 00007128) ( CAS), 13 (00007258) ( UCR) ; 28 Nov 2001 – 06 Dec 2001, M. species is very similar to G. quadocris , but is dis- Irwin, R. Harin’Hala , 13 (00007200) (SU) ; 02 Jan tinguished from G. quadocris by the larger body 2002– 10 Jan 2002, M. Irwin, R. Harin’Hala, 13 size, slightly more elongate head, more broadly (00007153) ( CAS) ; 10 Jan 2002 – 14 Jan 2002, M. Irwin, depressed postclypeus, labial segment II longer R. Harin’Hala , 13 (00007172) ( CAS) ; 12 Feb 2002 –19 than III, meso- and metasterna distinctly sepa- Feb 2002, M. Irwin, R. Harin’Hala , 13 (00007236) rated by a distinct transverse suture, and shape

(CAS); 31 Mar 2002 – 07 Apr 2002, M. Irwin, R. Harin’Hala , 13 (00045552) ( CAS) ; 07 Apr 2002 –14 Apr of the pygophore process in caudal view. 2002, M. Irwin, R. Harin’Hala, 23 (00044888, DESCRIPTION: MALE: Body length: 7.17 mm 00045425) ( AMNH) ; 14 Apr 2002 – 23 Apr 2002, M. (holotype), 6.80–7.61 mm. COLORATION: Dark Irwin, R. Harin’Hala, 33 (00044909, 00044949, brown with antennal segment V apically; anten- 00044951) ( USNM), 23 (00044954, 00045166) nal segments VI – VIII ; posterior margin of pro- (MNHN); 28 Apr 2002 – 05 May 2002, M. Irwin, R. Harin’Hala , 13 (00007184) ( CAS) ; 05 May 2002 –13 notum; corium basally; coxae; trochanters; basal May 2002, M. Irwin, R. Harin’Hala , 13 (00007186) half of femora ; anterior areas of laterotergites; (CAS); 13 Jun 2002 – 23 Jun 2002, M. Irwin, R. sternites medially, except pygophore ; and antero- Harin’Hala, 13 (00044802) ( CAS) ; 04 Jul 2002 –14 Jul laterally on sternites II – VI pale. Remainder of 2002, M. Irwin, R. Harin’Hala, 13 (00044895) ( CAS) ; corium and basal wing veins brown. VESTI- 24 Jul 2002 – 04 Aug 2002, M. Irwin, R. Harin’Hala , 13 TURE: Sparse ; other features as in generic

(00007219) (BMNH). Parc National Ranomafana, Vohiparara, at broken bridge, 21.22616 ° S 47.36983 ° E, description. STRUCTURE: HEAD: Ovoid in lat- 1100 m, 02 Jan 2002 – 10 Jan 2002, M. Irwin, R. eral view (pl. 13F) ; longer than wide in dorsal Harin’Hala, 13 (00007188) ( CAS) ; 19 Feb 2002 –26 Feb view; anteocular region as long as postocular (pl. 2002, M. Irwin, R. Harin’Hala , 13 (00007257) (SU); 04 13C) ; gula flat, conforming to rounded shape of Jun 2002– 14 Jun 2002, M. Irwin, R. Harin’Hala , 13 head (pl. 13C) ; postclypeus with shallow, broad (00044860) ( UCR). Parc National Ranomafana , radio medial longitudinal depression to middle of

tower at forest edge, 21.251 ° S 47.40716 ° E, 1130 m, 14 Jan 2002 – 21 Jan 2002, M. Irwin, R. Harin’Hala , 23 interocular area (pl. 13B) ; synthlipsis about 1.5 (00044882, 00045572) (BMNH); 21 Jan 2002 –28 Jan times width of eye; interocular sulcus posterior to 2002, M. Irwin, R. Harin’Hala , 13 (00045651) (SU). hind margin of eye ; ocelli small, located on shal- Parc National Ranomafana, radio tower at forest edge, low median tubercle (pl. 13I); eye about one- 21.25083 ° S 47.40716 ° E, 1130 m, 18 Mar 2006 –30 Mar fourth of head length, not reaching dorsal and

2006, M. Irwin, R. Harin’Hala, 13 (00007224) ( CAS). Ranomafana JIRAMA water works, 21.2485 ° S ventral head surfaces ; antenna inserted sublater- 47.45216 ° E, 690 m, 06 Dec 2001 – 15 Dec 2001, M. ally on head; pedicel about as long as scape; Irwin, R. Harin’Hala, 13 (00007166) ( CAS), 13 antenna 8-segmented ; labium stout; labial seg- (00007203) (AMNH); 21 Dec 2001 – 24 Dec 2001, M. ment III shorter than II, ventrally convex (pl. Irwin , R. Harin’Hala , 13 (00006465) ( UCR); 21 Jan 13C). THORAX: Anterior pronotal lobe with very 2002– 28 Jan 2002, M. Irwin, R. Harin’Hala, 13 small anterolateral projections (pl. 15B) ; pronotal

(00045348) (CAS). Unknown: 43 (00007117, 00007180, 00007181, 00007261) (CAS). transverse furrow continuous; lateral depressions on posterior pronotal lobe smooth; scutellar api-

cal projections dorsally directed (pl. 15C); meso-

Gibbosella vangocris , new species and metasterna separated by distinct transverse

Plates 1, 5, 8, 19B, 20C, 21B, 22A; map 3 suture (pl. 15F); mesosternum with medial and DIAGNOSIS: Males recognized among other paramedial longitudinal depressions; metaster- Gibbosella species by the relatively larger body num medially longitudinally depressed; legs slensize, slightly broader postclypeal depression, der; fossula spongiosa on mid tibia; tarsomeres I synthlipsis 1.5 times the width of an eye, labial and II combined subequal to III; hemelytron sur-

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 55

passing abdominal apex; distal part of R present Irwin, R. Harin’Hala , 63 (00048028–00048033) ( CAS) ; but not forming R +M cell (pl. 17B, C). ABDO- 15 Dec 2001 – 21 Dec 2001, M. Irwin, R. Harin’Hala, 13 MEN: Apex rounded ; sternites medially convex, (00045586) (SU); 24 Dec 2001 – 02 Jan 2002, M. Irwin, R. Harin’Hala , 33 (00007093, 00044929, 00045720)

with shallow medial depression on posterior half (USNM); 19 Feb 2002 – 26 Feb 2002, M. Irwin, R. of segment VI, intersegmental sutures carinulate Harin’Hala , 13 (00007016) ( UCR) ; 31 Mar 2002 –07 between II – VI ; pygophore (pls. 19B, 20C) process Apr 2002, M. Irwin, R. Harin’Hala, 53 (00048052– subtriangular basally and apically flattened and 00048056) ( CAS) ; 07 Apr 2002 – 14 Apr 2002, M. Irwin, curved in lateral view, spadelike in caudal view, R. Harin’Hala , 13 (00048034) (SU), 63 (00048036– 00048041) ( CAS) ; 14 Apr 2002 – 23 Apr 2002, M. Irwin ,

apex rounded; BPE as long as basal plate (pl.

R. Harin’Hala, 13 (00045342) (CAS); 05 May 2002 –13

22A); endosomal struts reaching posterior mar- May 2002, M. Irwin, R. Harin’Hala, 23 (00006985, gin of DPS, with posterior ventral process (pl. 00007013) (AMNH); 13 May 2002 – 23 May 2002, M. 22G), fused anterior to fusion with DPS; area of Irwin, R. Harin’Hala, 13 (00007027) (CAS); 13 Jun endosomal struts–DPS fusion ovate (pl. 21B). 2002– 23 Jun 2002, M. Irwin, R. Harin’Hala, 13

ETYMOLOGY: The species epithet is a noun in (00045716) (CAS); 04 Jul 2002 – 14 Jul 2002, M. Irwin, R. Harin’Hala, 13 (00007273) ( BMNH) ; 14 Jul 2002 –

apposition and is named for the spadelike 24 Jul 2002, M. Irwin, R. Harin’Hala, 13 (00045172) median pygophore process. (BMNH); 24 Jul 2002 – 04 Aug 2002, M. Irwin, R.

DISTRIBUTION: Marojejy National Park in the Harin’Hala, 23 (00007032, 00007280) ( USNM) ; 15 Antsiranana province, Ranomafana National May 2003– 28 May 2003, M. Irwin, R. Harin’Hala, 13 Park in the Fianarantsoa province, and Berenty (00006993) ( UCR). Parc National Ranomafana, Vohiparara, at broken bridge, 21.22616 ° S 47.36983 ° E, 1100 m ,

Special Reserve in the Toliara province (map 3).

28 Nov 2001 – 06 Dec 2001, M. Irwin, R. Harin’Hala, 43

DISCUSSION: The coloration is slightly variable (00044845, 00044870, 00045020, 00045230) (BMNH), with the posterior pronotal lobe, corium, and 33 (00045406, 00045409, 00045676) (CAS), 13 basal wing veins ranging from pale to brown. (00045688) (UCR); 02 Jan 2002 – 10 Jan 2002, M. Irwin,

HOLOTYPE: Male : Madagascar: Fianarant- R. Harin’Hala , 23 (00006999, 00007023) ( AMNH); 28 soa: Parc National Ranomafana , Belle Vue at Jan 2002– 04 Feb 2002, M. Irwin, R. Harin’Hala, 43 (00048042–00048045) ( MNHN), 63 (00048046–

Talatakely, 21.2665 ° S 47.42016 ° E, 1020 m, 16 Oct

00048051) ( CAS); 19 Mar 2002 – 26 Mar 2002, M. Irwin,

2001– 08 Nov 2001, M. Irwin, R. Harin’Hala R. Harin’Hala , 23 (00045199, 00045384) (SU) ; 26 Mar (00006995) ( CAS). 2002– 31 Mar 2002, M. Irwin, R. Harin’Hala , 13

PARATYPES: Madagascar: Antsiranana: Marojejy (00007040) ( CAS) ; 08 Apr 2002 – 15 Apr 2002, M. Irwin, National Park, 5 km W Manantenina village, Camp R. Harin’Hala , 23 (00007042, 00007097) ( CAS) ; 22 Apr Mantella, 14.43816 ° S 49.774 ° E, 490 m, 25 Feb 2005 –04 2002– 29 Apr 2002, M. Irwin, R. Harin’Hala, 23 Mar 2005, M. Irwin, R. Harin’Hala , 13 (00007275) (00007272, 00007278) ( CAS) ; 06 May 2002 –15 May ( CAS) . Fianarantsoa: Parc National Ranomafana, Bev- 2002, M. Irwin, R. Harin’Hala , 13 (00007015) ( UCR) ; aohazo-Ranomana, 21.2 ° S 47.48333 ° E, 970 m, Nov 04 Jun 2002 – 14 Jun 2002, M. Irwin, R. Harin’Hala, 23 2000, M. Irwin, R. Harin’Hala , 13 (00044919) ( CAS). (00007116, 00007119) ( USNM) ; 15 Jul 2002 –25 Jul Parc National Ranomafana, Belle Vue at Talatakely, 2002, M. Irwin, R. Harin’Hala, 23 (00045313, 21.2665 ° S 47.42016 ° E, 1020 m, 16 Oct 2001 –08 Nov 00045578) ( AMNH). Parc National Ranomafana, radio 2001, M. Irwin, R. Harin’Hala, 33 (00006984, tower at forest edge, 21.251 ° S 47.40716 ° E, 1130 m, 06 00007000, 00007001) ( USNM), 133 (00007005, Dec 2001– 15 Dec 2001, M. Irwin, R. Harin’Hala , 13 00007014, 00007022, 00007029, 00007030, 00007033, (00007011) ( MNHN), 13 (00007213) ( CAS) ; 15 Dec 00007055, 00007057, 00007063, 00007070, 00007071, 2001– 21 Dec 2001, M. Irwin, R. Harin’Hala, 13 00007077, 00007092) ( CAS), 43 (00007111, 00007114, (00045588) ( UCR); 21 Dec 2001 – 24 Dec 2001, M. 00007126, 00007254) ( MNHN), 33 (00007268, Irwin, R. Harin’Hala , 23 (00007019, 00045037) ( SU); 00007270, 00048227) ( AMNH) ; 08 Nov 2001 –15 Nov 14 Jan 2002 – 21 Jan 2002, M. Irwin, R. Harin’Hala, 23 2001, M. Irwin, R. Harin’Hala, 33 (00006085, (00045516, 00045554) ( AMNH); 19 Feb 2002 –26 Feb 00007006, 00007017) ( MNHN), 43 (00007018, 2002, M. Irwin, R. Harin’Hala , 13 (00007008) ( CAS); 00007044, 00007144, 00007274) ( BMNH) ; 15 Nov 12 Mar 2002 – 19 Mar 2002, M. Irwin, R. Harin’Hala, 2001– 22 Nov 2001, M. Irwin, R. Harin’Hala , 13 13 (00006994) ( CAS) ; 23 Apr 2002 – 30 Apr 2002, M. (00045570) ( UCR) ; 22 Nov 2001 – 28 Nov 2001, M. Irwin, R. Harin’Hala , 13 (00007187) ( SU) ; 14 Jun

56 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

2002– 24 Jun 2002, M. Irwin, R. Harin’Hala, 23 (Maldonado, 1990). Within Madagascar, the two

(00006997, 00007007) ( BMNH); 24 Jun 2002 –05 Jul species of Glymmatophora are known from the

2002, M. Irwin, R. Harin’Hala, 13 (00007105) ( UCR); Toliara and Fianarantsoa provinces. Specimens

15 Jul 2002 – 25 Jul 2002, M. Irwin, R. Harin’Hala, 23 have been collected in spiny forest thicket,

(00007004, 00007025) ( USNM); 18 Feb 2003 –27 Feb

2003, M. Irwin, R. Harin’Hala, 13 (00045078) degraded grassland, and in the Antsarongaza dry

( AMNH); 20 Mar 2003 – 03 Apr 2003, M. Irwin, R. forest between 18–825 m elevation.

Harin’Hala, 13 (00007121) (SU); 06 Jul 2003 –17 Jul DISCUSSION: The largely Afrotropical genus

2003, M. Irwin, R. Harin’Hala, 13 (00007269) ( CAS); Glymmatophora was described by Carl Stål in

27 Aug 2003 – 07 Sep 2003, M. Irwin, R. Harin’Hala, 13 1853. In 1914, Géza Horváth described three

(00007021) ( UCR); 28 Sep 2003 – 08 Oct 2003, M.

Irwin, R. Harin’Hala, 23 (00007002, 00007233) ( CAS); subgenera: ( Cyclosandalus ), ( Glymmatophora ),

02 Apr 2004 – 15 Apr 2004, M. Irwin, R. Harin’Hala, 13 and ( Haematorrhophus ), the last of which is cur-

(00007003) ( USNM); 11 Jun 2004 – 24 Jun 2004, M. rently treated as a distinct genus. The former two

Irwin, R. Harin’Hala, 13 (00045507) ( CAS); 24 Jun subgenera were primarily diagnosed based on

2004– 08 Jul 2004, M. Irwin, R. Harin’Hala, 13 the presence (subgenus Glymmatophora ) or

(00007169) ( AMNH); 23 Sep 2004 – 07 Oct 2004, M. absence (subgenus Cyclosandalus ) of ocelli in the

Irwin, R. Harin’Hala, 13 (00045660) (SU); 11 Dec

2004– 18 Dec 2004, M. Irwin, R. Harin’Hala, 13 female and the size and shape of the fossula

(00045625) ( UCR); 02 Feb 2005 – 17 Feb 2005, M. Irwin, spongiosa (large and oblong in subgenus Glym-

R. Harin’Hala, 13 (00045563) ( CAS); 27 Feb 2005 –13 matophora; small and round in subgenus Cyclos-

Mar 2005, M. Irwin, R. Harin’Hala, 13 (00007069) andalus). Prior to this monograph, one species of

( UCR); 25 Jul 2006 – 09 Aug 2006, M. Irwin, R. Glymmatophora was known from Madagascar ,

Harin’Hala, 13 (00045371) ( CAS). Ranomafana

JIRAMA water works, 21.2485 ° S 47.45216 ° E, 690 m, 22 G. (Cyclosandalus) crassipes . A new Madagascan

Nov 2001– 28 Nov 2001, M. Irwin, R. Harin’Hala, 13 species of Glymmatophora is here described, G.

(00006124) ( UCR); 06 Dec 2001 – 15 Dec 2001, M. (Glymmatophora) carolae .

Irwin, R. Harin’Hala, 43 (00006982, 00006988,

00006991, 00006996) ( CAS), 23 (00007020, 00007031) IDENTIFICATION KEY TO THE MALES AND

(SU); 21 Dec 2001 – 24 Dec 2001, M. Irwin, R. FEMALES OF MADAGASCAN SPECIES OF

Harin’Hala, 13 (00007276) ( CAS); 24 Dec 2001 –02 Jan

2002, M. Irwin, R. Harin’Hala, 13 (00006986) ( UCR); Glymmatophora

21 Jan 2002 – 28 Jan 2002, M. Irwin, R. Harin’Hala, 13 1. Males only known; macropterous; ocelli pres-

(00044847) (SU), 13 (00045687) ( CAS). Toliara: ent... (Glymmatophora) carolae , new species

Berenty Special Reserve, 8 km NW Amboasary, – Males and females apterous; ocelli absent (pl.

25.021 ° S 46.3055 ° E, 35 m, 29 Aug 2004 – 08 Sep 2004, 13D)..... (Cyclosandalus) crassipes Horváth

M. Irwin, F. Parker, R. Harin’Hala, 13 (00007197)

( UCR), 13 (00007248) ( CAS), 13 (00007250) (SU).

Unknown: 123 (00006983, 00006987, 00006989, Glymmatophora (Glymmatophora) carolae ,

00006990, 00006992, 00007010, 00007024, 00007026, new species

00007221, 00007271, 00007277, 00007279) ( CAS). Plates 1, 5, 8, 16C, 17B, 20D, 21C; map 4

DIAGNOSIS: Males are recognized among

Glymmatophora Stål, 1853 other Glymmatophora species by the very dis-

Plates 1, 2, 5, 8, 12, 13A, 13D, 15E, 16B, 16C, tinctive coloration (mostly red with some black

16E, 17B, 19C, 20D, 21C, 21D, 22B, 23C, 23D, markings), anteriorly concave postclypeus, fore- 23F; map 4 femur with anterior subapical and medial protu-

Glymmatophora Stål, 1853: 261 . berances and macropterous condition. This species is very similar to the Afrotropical G.

TYPE SPECIES: Glymmatophora submetallica lothei Villiers, 1948 , from which it differs by the

Stål, 1853. slightly smaller body size, black antenna, red

DISTRIBUTION: Thirty-five described species abdomen with black markings (i.e., anterior

of Glymmatophora occur throughout Africa band on laterotergites IV and V, laterotergite VI

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 57

except posterolaterally, anterior half of lateroter-

gite VII, lateral margin of abdominal sternite VI,

and anterolateral margin of sternite VII), more

medially concaved anterior pronotal margin, and

forefemur with only two protuberances.

DESCRIPTION: MALE: Macropterous, medium

body size (length: 16.91 mm, holotype; 14.90–

16.91 mm). COLORATION: Red with antenna,

hemelytron except corium basally and anteriorly,

anterior band on laterotergites IV and V, lateroter-

gite VI except posterolaterally, anterior half of lat-

erotergite VII, lateral margin of abdominal sternite

VI, and anterolateral margin of sternite VII black.

VESTITURE: Glabrous, except antenna with

long, erect brown setae and tibiae with stout pale

brown setae that are denser near apex. STRUC-

TURE: HEAD: Subquadrate in lateral view (pl.

13D); wider than long in dorsal view; ventrally

flat; anteocular region as long as postocular; gula

swollen ventrolaterally, slightly produced beyond

ventral margin of head (pl. 13E); maxillary plate

not reaching dorsal clypeal surface (pl. 13E, I);

postclypeus concave anteriorly with shallow, very

narrow medial longitudinal depression to middle

of interocular area; synthlipsis (pl. 13A) about two

times width of eye; interocular sulcus near hind

margin of eye (pl. 13B); ocelli small, located on

distinct median tubercle (pl. 13H), separated by MAP 4. Localities of Glymmatophora species and more than diameter of ocellus; eye about one- Maraenaspis bidens .

third head length, not reaching dorsal and ventral

head surfaces; antennal shield concealing antennal 15E); mesosternum with medial and paramedial insertion in lateral view (pl. 13D); antenna longitudinal depressions (pl. 15F); metasternum inserted sublaterally on head; pedicel about one- medially longitudinally depressed (pl. 15F); MGE fourth longer than scape, slightly curved. THO- with shallow meshlike cuticle that does not extend RAX: Pronotum wider than long, anterior margin dorsally in lateral view (pl. 16B, C); forefemur moderately concave, smooth; anterior pronotal with anterior subapical and medial protuberances lobe shorter than posterior lobe (pl. 15B), more (pl. 16E), base ventrally carinate; mid and hind than half as wide as posterior lobe, without dis- femora with anterior and posterior subapical and tinct anterolateral projections; pronotal longitudi- medial protuberances; foretibia with ventral spine nal furrow reaching anterior but not posterior basal to fossula spongiosa; hemelytron (pl. 17B) margin, weakly foveate posteriorly (pl. 15B); pro- surpassing abdominal tergite VII but not reaching notal transverse furrow divided by paramedian apex; corium well developed; distal part of R preslongitudinal ridges (pl. 15A, B); lateral depres- ent but not forming R+M cell; proximal parts of sions of posterior pronotal lobe distinct (pl. 15B), M and Cu fused; distal part of M extending weakly transversely striated; meso- and metasterna beyond apical junction of M+Cu; base of M+Cu not completely separated by transverse suture (pl. cell shorter than or as wide as Cu+1A cell. ABDO-

58 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

MEN: Apex rounded; dorsal laterotergite II forms of G. lothei , but the presence of two protuexpanded; intersegmental sutures carinulate berances on the forefemur distinguishes G. between sternites II–VI and laterally between VI crassipes from G. lothei .

and VII; sternites shallowly depressed medially REDESCRIPTION: MALE: Apterous , medium (pl. 18C) ; spiracles ovoid; pygophore process dor- body size (length: 15.52–19.60 mm). COLORsally knoblike in lateral view (pl. 20D), subtrian- ATION: Black with head, labium, scape, pedicel gular in caudal view (pl. 19C), directed basally, pronotum except pronotal longitudinal dorsoposteriad, not surpassing posterior margin furrow, propleuron except dorsal spot and stripe of pygophore, apex rounded ; BPE shorter than on anterior lobe, meso- and metapleura dorsad of basal plate; DPS apex concave medially (pl. 21C); coxae, meso- and metasterna laterally, legs, latendosomal struts reaching posterior margin of erotergite II, laterotergite III and IV posteriorly, DPS (pl. 21C), ventrally obscured by semiextend sternites II–VI transversely basally, sternite VII endosoma; area of endosomal struts–DPS fusion posteriorly, and pygophore red. VESTITURE: elongate subquadrate; endosoma medially sclero- Glabrous, except antenna with short, semierect to tized posteriorly and sclerotization extending erect brown setae and tibiae with stout, pale anterolaterally (pl. 21C). brown setae that are denser near apex. STRUC- ETYMOLOGY: This species epithet is a noun in TURE: HEAD (pl. 13A, D): Subquadrate in lateral the genitive case and is named in memoriam of view ; longer than wide in dorsal view; ventrally the senior author’s mother, Carol Forthman. flat; anteocular region longer than postocular; DISTRIBUTION: Antsarongaza dry forest in the gula swollen ventrolaterally, not produced beyond Toliara province (map 4). ventral margin of head, carinate ; maxillary plate DISCUSSION: Female specimens are unknown. not reaching dorsal clypeal surface (pl. 13E, I) ; We assign this species to the subgenus Glymma- postclypeus flat ; synthlipsis about 2.5 times width tophora based on the features mentioned in the of eye; interocular sulcus near hind margin of eye generic discussion. (pl. 13B); eye about one-fourth of head length, HOLOTYPE: Male : Madagascar: Toliara: not reaching dorsal and ventral head surfaces ; Beroboka village , 45 km NE Morondava, 19.9775 ° S antennal shield concealing antennal insertion in 44.82483 ° E, 131 m, 17 Nov 2008 – 25 Nov 2008, M. lateral view ; antenna inserted sublaterally on Irwin, R. Harin’Hala (00044889) ( CAS). head ; pedicel slightly longer than scape, slightly PARATYPES: Madagascar: Toliara: Bekily, curved. THORAX (pl. 15E): Pronotum as long as 24.22802 ° S 45.30858 ° E, 388 m, Oct 1936, A. Seyrig, 23 wide, anterior margin slightly concave, smooth; ( MNHN) ; Beroboka village , 45 km NE Morondava ,

anterior pronotal lobe longer than and as wide as 19.9775 ° S 44.82483 ° E, 131 m, 05 Feb 2009 –13 Feb

2009, M. Irwin, R. Harin’Hala, 13 (00045479) (CAS). posterior lobe (pl. 15A), without distinct antero-

lateral projections; pronotal longitudinal furrow Glymmatophora (Cyclosandalus) crassipes not reaching anterior and posterior margins,

Horváth, 1914 reduced to deep medial depression near posterior Plates 1, 2, 5, 8, 12, 13A, 13D, 15E, 16B, 16E, margin of anterior lobe (pl. 15A); pronotal trans- 19C, 21D, 22B, 23C, 23D, 23F; map 4 verse furrow weakly to strongly divided by para- Glymmatophora (Cyclosandalus) crassipes Hor- median longitudinal ridges (pl. 15A); lateral

depressions of posterior pronotal lobe obsolete; váth, 1914: 128.

meso- and metasterna not completely separated REVISED DIAGNOSIS: Males and females are by transverse suture; mesosternum with medial recognized among other species in this genus by and paramedial longitudinal depressions; metathe apterous condition, lack of ocelli in both sternum medially longitudinally depressed; MGE sexes, and fossula spongiosa on foretibia small with shallow meshlike cuticle that does not and round. This species is similar to apterous extend dorsally in lateral view (pl. 16B, C); tro-

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 59

chanters and fore- and mid femora basally with from “ P. Majanga ” in the Mahajanga province. patches of small papillae (pl. 16E); forefemur with It is not clear if “ P. Majanga ” is referring to a anterior subapical and medial protuberances (pl. park within the Mahajanga province or the 16E); mid and hind femora with anterior and town Mahajanga. posterior subapical and medial protuberances; DISCUSSION: A few specimens without or with foretibia with ventral spine basal to fossula spon- reduced red coloration on the scutellum, laterogiosa. ABDOMEN: Apex rounded; dorsal lateroter- tergites, and sternites were examined and are gite II slightly expanded; intersegmental sutures treated as conspecific with G. crassipes based on carinulate between sternites II – VI and laterally corresponding external and internal morpholbetween VI and VII; sternites medially flat; spir- ogy. Specimens of both color morphs were colacles ovoid; pygophore (pls. 19C, 20D) process lected in close proximity. dorsally knoblike in lateral view, subtriangular in HOLOTYPE: Female : Madagascar: Toliara: caudal view, directed dorsoposteriad, not sur- Tsivory , 24.0684 ° S 46.07471 ° E, 431 m, 1906 Faupassing posterior margin of pygophore, apex chère ( HNHM). rounded; BPE about as long as basal plate (pl. OTHER MATERIAL EXAMINED: Madagascar: Anta- 22B); DPS apex rounded (pl. 21D); endosomal nanarivo: Amboasary ( Imerina ), 1899, G. Grandidier, struts reaching posterior margin of DPS, with 1♀ ( MNHN). Fianarantsoa: Ikongo , 21.99483 ° S anterior and posterior ventral processes (pl. 22G); 47.37065 ° E, 536 m, 1902, G. Grandidier, 13 ( MNHN). area of endosomal struts–DPS fusion elongate Mahajanga: P. Majanga, Nov 1965, Malzy 13 ( MNHN). Toliara: 18 km NW Betroka, 23.16333 ° S 45.96861 ° E, 825 subquadrate (pl. 21D); endosoma medially scler- m, 04 Dec 1994 – 09 Dec 1994, M.A. Ivie and D.A. Polotized posteriorly and extends anterolaterally (pl. lock, 13 (00120258) ( MTEC). Ankazoabo, 21.50348 ° S 21D). FEMALE: Body length: 16.50 mm (holo- 45.20956 ° E, 278 m, 1902, J. Bastard, 1♀ ( MNHN); 1920, type; from original description), 16.50–23.90 G. Le Barbier, 13 ( MNHN); no date provided, 1♀ mm. Similar to males, but differ in the following ( MNHN). Antaloba, no date provided, 1♀ ( MNHN). Bas Mangoky, Agriculture Station , no date provided, 13, characters: COLORATION: Red on prothorax, 1♀ ( MNHN). Behara, 24.95347 ° S 46.38593 ° E, 49 m, 10 mesopleuron, metapleuron dorsad of coxa, lateral Aug 1918– 11 Aug 1918, A. R., 1♀ ( MNHN). Bekily, metanotal spots, scutellum laterally, meso- and 24.22802 ° S 45.30858 ° E, 388 m, Mar 1936, A. Seyrig, 13 metasterna, laterotergite II, posterolateral margin ( MNHN). Berenty Reserve , W of Fort Dauphin, of laterotergites III – VII, and sternite VII postero- 25.00564 ° S 46.30278 ° E, 09 Nov 2008 – 24 Nov 2008, J. R. medially. STRUCTURE: THORAX: Fossula spon- Cryan and G. Svenson, 23 (00088087, 00088088) ( UCR). Betroka , Andalamby, 23.26833 ° S 46.10444 ° E, giosa larger. ABDOMEN: External genitalia as in 822 m, Jan 1958, J. Elie, 1♀ ( MNHN). Country Mahaplate 23C, D; bursa copulatrix membranous, with faly, 1906, J. Bastard, 13 ( MNHN). Lambomakandro, lateral lobes (pl. 23F); vermiform gland and lat- 22.69831 ° S 44.70099 ° E, 526 m, 1936, R. Catala 1♀ eral spermathecae missing. ( MNHN). Mahafaly Plateau, 6.2 km 74° ENE Itampolo, DISTRIBUTION: Most specimens were col- 24.65361 ° S 43.99666 ° E, 80 m, 21 Feb 2002 – 25 Feb 2002, Fisher et al., 1♀ (00007050) ( CAS). Mahafaly Plateau, lected in the Fianarantsoa and Toliara provinces Ankalirano, 25.45 ° S 45.71667 ° E, 18 m, 17 Jan 1974, P. and were successfully georeferenced (map 4). Viette & A. Peyrieras, 1♀ ( MNHN). On plateau, Some could not be georeferenced beyond prov- 23.00283 ° S 43.70366 ° E, 30 m, 26 Feb 2003 – 08 May 2003, ince based on locality data, with most of them Frontier Wilderness Project, CAS, 1♀ (00006372) ( CAS). collected in the Toliara province. One specimen Onilahy River , 1922, G. Petit, 1♀ ( MNHN). Ranobe, was associated with historical locality data that 23.00943 ° S 43.60934 ° E, 20 m, 1905, F. Geay, 2♀ ( MNHN). Réserve Spéciale de Cap Sainte Marie, 12.3 could not be georeferenced (i.e., “Amboasary km 262° W Marovato, 25.58166 ° S 45.16833 ° E, 200 m, 11 [Imerina]”). Imerina refers to a kingdom that Feb 2002– 15 Feb 2002, Fisher et al., 13 (00044857) once existed and likely is part of modern-day ( CAS). Zombitse-Vohibasia National Park, Eastern Antananarivo province. A second specimen Sakaraha, Matsabory, 22.8417 ° S 44.68184 ° E, 640 m, 13 could not be georeferenced but was collected Feb 1974, P. Viette and A. Peyrieras, 13 ( MNHN).

60 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

Maraenaspis Karsch, 1892 pale to light brown. VESTITURE: Glabrous, Plates 2, 5, 8, 13E, 15A, 19D, 20E, 21E, 22C; except antenna with short, semierect to erect map 4 pale to brown setae and tibiae with stout pale

orange-brown setae that are denser near apex. Maraenaspis Karsch, 1892 , 18: 134.

STRUCTURE: HEAD (pl. 13E): Ovoid in lateral

TYPE SPECIES: Maraenaspis typhlops Karsch , view; longer than wide in dorsal view; ventrally 1892. flat; anteocular region longer than postocular;

DISTRIBUTION: Thirty-eight described species gula swollen ventrolaterally, produced beyond of Maraenaspis occur throughout the Afrotropics ventral margin of head, carinate; maxillary plate (Maldonado, 1990). Specimens of the only known not reaching dorsal clypeal surface; postclypeus Madagascan species Maraenaspis have been col- flat; antennal shield not concealing antennal lected from Nosy Be Island off the northwest insertion in lateral view; synthlipsis about two coast, as well as the Antsiranana, Mahajanga, and times width of eye; interocular sulcus posterior Toamasina provinces. Records are from tropical to hind margin of eye; eye about one-fourth of dry forest between 8–135 m elevation. head length, not reaching dorsal and ventral

DISCUSSION: Maraenaspis was described by head surfaces; antenna inserted sublaterally on Ferdinand Karsch in 1892. Maraenaspis bidens head; pedicel about one-fourth longer than (Reuter), which was originally described from an scape, slightly curved. THORAX (pl. 15A): Proapterous female, is the only species of Maraenas- notum as long as wide, smooth; anterior pronopis known from Madagascar. The male of M. tal lobe with large anterolateral projections; bidens is described here for the first time. pronotal longitudinal furrow reaching anterior

but not posterior margins, deeply depressed Maraenaspis bidens (Reuter), 1887 near posterior margin of anterior lobe; pronotal Plates 2, 5, 8, 13E, 15A, 19D, 20E, 21E, 22C; transverse furrow divided by paramedian longimap 4 tudinal ridges; lateral depressions of posterior Cleptria bidens Reuter, 1887: 108 . pronotal lobe obsolete; meso- and metasterna

not completely separated by transverse suture Maraenaspis bidens: Villiers, 1945: 199 .

(pl. 15E); mesosternum with medial and para-

REVISED DIAGNOSIS: Males and females are medial longitudinal depressions; metasternum recognized among other species of Maraenaspis medially longitudinally depressed; MGE with by a combination of the following characters: the shallowly depressed meshlike cuticle that does large body size, coloration, anterior pronotal not extend dorsally in lateral view (pl. 16B, C); lobe with very large anterolateral projections, trochanters and fore- and mid femora basally apterous condition, ovoid abdomen that is api- with small papillae (pl. 16E). ABDOMEN: Apex cally notched, and sternites convex. This species medially notched; dorsal laterotergite II is similar to M. neglecta Villiers, 1948 , but is dis- expanded; intersegmental sutures carinulate tinguished by the slightly larger body size, blu- between sternites II and III and laterally between ish-black coloration on the head and legs, and III–VII; sternites medially flat; spiracles elliptivery strong anterolateral protuberances on the cal; pygophore (pls. 19D, 20E) process hooklike anterior pronotal lobe. in lateral view, subtriangular in caudal view,

REDESCRIPTION: MALE: Apterous; large directed dorsoposteriad, not surpassing postebody size (length: 21.63–22.08 mm). COLOR- rior margin of pygophore, apex rounded; BPE

ATION: Bluish-black suffused with violet sheen shorter than basal plate; DPS apex rounded (pl. on abdominal tergites. Antennal segments III, 21E); endosomal struts reaching posterior mar- IV, base of V, and apex of VIII brown and anten- gin of DPS, ventrally obscured by semiextended nal segments V apically, VI, VII, and VIII basally endosoma; area of endosomal struts–DPS fusion

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 61

circular (pl. 21E); endosoma with platelike scler- m, 02 Apr 2001 – 08 Apr 2001, Fisher et al., 13 (00045017) ite on posterior margin (pl. 21E). FEMALE: ( CAS). Toamasina: Alahakato Forest, 1888, Edouard Body length: 21.5 mm (holotype; original Perrot, 1♀ ( MNHN). Antanambe, 16.43333 ° S 49.85 ° E, 8

m, 1898, A. Mocquerys, 1♀ (MNHN). description), 21.30–23.00 mm; coloration and structure same as the male. DISTRIBUTION: Natural Reserve of Lokobe on Marojejycoris , new genus Nosy Be Island, and the Antsiranana, Mahajanga, Plates 2, 5, 6, 9, 13F, 19E, 20F, 21F, 22D; map 5 and Toamasina provinces (map 4). TYPE SPECIES: Marojejycoris notadichroa , new DISCUSSION: This species was described from species. a single female specimen collected (coll. Stumpff) DIAGNOSIS: Males are recognized by the small in “Loucoubé,” which is currently known as the body size, dull coloration, dense vestiture, Natural Reserve of Lokobe on Nosy Be Island. broadly depressed postclypeus, ocelli located on Putshkov and Putschkov’s (1986) Ectrichodiinae a shallow medial ocellar tubercle, large antennal catalog indicates that the holotype for M. bidens shield that does not conceal the antennal inseris deposited in the National Museum of Natural tion in lateral view, continuous transverse suture History, Paris, France, but the type could not be on the pronotum, MGE with deeply depressed located (Eric Guilbert, personal commun.). Cor- meshlike cuticle the extends dorsally in lateral respondence with other major European collec- view, distal part of M not extending beyond apitions did not offer any further leads, and we cal junction of M+Cu, dorsal laterotergite II not conclude that the type may be lost or placed expanded, and circular spiracles. Marojejycoris within undetermined material. Given the poten- species are similar to the Afrotropical genus Syntial that the holotype may be located among avecoris, but are distinguished by the denser vesundetermined material that we have not exam- titure, head longer than wide in dorsal view, ined, we refrain from designating a neotype. depressed postclypeus, presence of the antennal The male and female specimens examined shield, more medially excavated anterior margin match Reuter’s original description of the spe- of the pronotum, anterior pronotal lobe lacking cies, with the exception of the antennal color- carinae laterally, continuous transverse suture, ation. The original description states that the last meso- and metasterna separated by a distinct four antennal segments in the female are “stra- transverse suture, MGE extending dorsally in latminous” with the “base of the fifth yellowish.” In eral view, unarmed legs, proximal part of M and the females examined, the distal antennal seg- Cu veins not fused, distal parts of M and Cu not ments were missing; in the males, antennal seg- fused, and dorsal laterotergites uniform in color ments III, IV, base of V, and apex of VIII are and not expanded. brown, while segments V apically, VI, VII, and DESCRIPTION: MALE: Macropterous, small VIII basally are pale to light brown. This slight body size. COLORATION: Orange-brownish or difference is considered to be variation within brown with pale markings. VESTITURE: Dense, the species or between the sexes. semierect to erect, long pale to brown setae on HOLOTYPE: Female: Madagascar: Toliara: head, thorax, legs, corium of hemelytra, and Nosy Be, Lokobe Natural Reserve, 13.39917 ° S abdomen; setae on tibiae stouter and denser near 48.31833 ° E, 204 m (see Discussion regarding apex. STRUCTURE: HEAD (pl. 13F): Ovoid or depository). subtriangular, longer than wide in dorsal view, OTHER MATERIAL EXAMINED: Madagascar: Antsira- shorter than pronotum; anteocular region as nana: Forêt d’Anabohazo, 21.6 km 247° WSW Maroman- long as postocular; ventrally with shallow anterodia, 14.30888 ° S 47.91444 ° E, 120 m, 11 Mar 2001 –16 Mar

medial depression; clypeal apex not elevated 2001, Fisher et al., (00044913) ( CAS). Mahajanga: Parc National d’Ankarafantsika, Forêt de Tsimaloto , 18.3 km relative to labrum ; maxillary plate not reaching 46° NE de Tsaramandroso , 16.22805 ° S 46.14361 ° E, 135 dorsal surface of clypeus (pl. 13E, I) ; postclypeus

62 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 400

broadly depressed (pl. 13B); postocular broad in sternites longitudinally depressed medially (pl. dorsal view (pl. 13B); ocelli present, located on 18C), intersegmental sutures carinulate; spiracles shallow medial tubercle, separated by less than circular; pygophore (pls. 19E, 20F) process flatdiameter of ocellus; distinct constriction between tened in lateral view, broadly rodlike in caudal postocular and neck; gula moderately swollen view, directed dorsoposteriad, not surpassing ventrolaterally, not distinctly produced beyond posterior margin of pygophore, apex rounded; ventral head margin; antennal shield not con- DPS apex rounded (pl. 21F); endosomal struts cealing antennal insertion in lateral view; reaching posterior margin of DPS, with anterior antenna inserted dorsally on head; scape sur- and posterior ventral processes (pl. 22G); endopassing clypeal apex; pedicel slightly curved; fla- soma sclerotized medially (pl. 21F).

gellum subdivided into BFLA and DFLA; BFLA ETYMOLOGY: The genus is a named after the divided into two pseudosegments; DFLA divided type locality of the type species, Marojejy National into two or three pseudosegments ; antenna thus Park, Madagascar. The gender is feminine.

appearing 6- or 7-segmented; labium relatively DISTRIBUTION: Species are known from the slender ; labial segment III shorter than II, ven- Antsiranana, Fianarantsoa, and Toamasina provtrally convex (appears dorsal when reposed). inces and in habitats between 10–1130 m elevation. THORAX: Pronotum wider than long, anterior Macrohabitats are described as mixed tropical formargin distinctly concave, collar distinct (pl. ests, low altitude rainforests and dense humid for- 15B) ; anterior pronotal lobe shorter than poste- ests, and forested limestone ridge.

rior lobe (pl. 15B), more than half as wide as DISCUSSION: Female specimens are unknown posterior lobe, without distinct anterolateral pro- for all species in this genus.

jections, relatively smooth; pronotal longitudinal

furrow reaching anterior but not posterior mar- IDENTIFICATION KEY TO THE MALES OF

gin of pronotum, weakly to strongly foveate pos- SPECIES OF MAROJEJYCORIS

teriorly (pl. 15B); pronotal transverse furrow 1. DFLA divided into two pseudosegments; continuous; lateral depressions on posterior pro- antenna thus appearing 6-segmented.... 2 notal lobe distinct (pl. 15B), transversely striated; – DFLA divided into three pseudosegments; scutellum with two moderately separated apical antenna thus appearing 7-segmented.... 3 processes, disc medially depressed; meso- and 2. Postclypeal depression extending to middle of metasterna separated by distinct transverse interocular area (pl. 13B); eye about onesuture (pl. 15F); mesosternum with medial and fourth head length in lateral view; scape, paramedial longitudinal depressions; metaster- pedicel, and antennal segments III–IV and num medially longitudinally depressed; MGE base of V dark brown, nearly black; corium with deeply depressed meshlike cuticle that orange........ auranticorium , new species extends dorsally in lateral view (pl. 16A); fossula – Postclypeal depression extending to interocular spongiosa on foretibia, absent on mid tibia in sulcus; eye about one-third head length in some species; tarsomeres I and II combined sub- lateral view; scape apically, antennal segequal to III; hemelytron surpassing abdominal ments II–IV, and base of V brown; corium apex; corium restricted to areas adjacent to basal brown.......... notadichroa , new species wing veins, with pterostigmalike appearance on 3. Fossula spongiosa absent or reduced on the anterodistal margin (pl. 17A, C); proximal parts mid tibia............................. 4 of M and Cu separate (pl. 17A, C); distal part of – Fossula spongiosa distinctly present on the M not extending beyond apical junction of mid tibia........ ranomafana , new species M+Cu; base of M+Cu cell shorter than or as 4. Synthlipsis about two times width of eye; eye wide as Cu+1A cell (pl. 17B, C). ABDOMEN: Apex not reaching dorsal and ventral head marrounded; dorsal laterotergites not expanded; gins................. francais , new species

2016 FORTHMAN ET AL.: MILLIPEDE ASSASSIN BUG FAUNA 63

– Synthlipsis about width of eye; eye reaching dorsal and ventral head margins (pl. 13F)................... brevifrons , new species