Notes on a collection of Myriapoda from Cuba

ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright copy 2013 Magnolia Press

Zootaxa 3626 (4) 477ndash498 wwwmapresscomzootaxa Article

httpdxdoiorg1011646zootaxa362644httpzoobankorgurnlsidzoobankorgpub24763A0D-5B5F-4DDB-9698-5D608738FE4D

The milliped family Platyrhacidae (Polydesmida Leptodesmidea) in the West Indies Proposal of Hoffmanorhacus n gen description and illustrations of males of Proaspis aitia Loomis 1941 redescription of Nannorrhacus luciae (Pocock 1894) hypotheses on origins and affinities and an updated New World familial distribution

ROWLAND M SHELLEY1 amp DANIELA MARTINEZ-TORRES2

1Research Laboratory North Carolina State Museum of Natural Sciences MSC 1626 Raleigh NC 27699-1626 USA E-mail rowlandshelleynaturalsciencesorg2Instituto de Ciencias NaturalesUniversidad Nacional de Colombia Edificio 425 Oficina 105 Bogotaacute Colombia E-mail martinezdanielagmailcom

Abstract

In the New World the milliped family Platyrhacidae (Polydesmida) is known or projected for Central America south of southeastern Nicaragua and the northern frac14 of South America with disjunct insular populations on Hispaniola (Haiti) Guadeloupe (Basse-Terre) and St Lucia Male near-topotypes enable redescription of Proaspis aitia Loomis 1941 possibly endemic to the western end of the southern Haitian peninsula The tibiotarsus of its biramous gonopodal telopodite bends strongly laterad and the medially directed solenomere arises at midlength proximal to the bend With a uniramous telopodite P sahlii Jeekel 1980 on Guadeloupe is not congeneric and Hoffmanorhacus n gen is erected to accommodate it Nannorrhacus luciae (Pocock 1894) on St Lucia is redescribed also with a biramous telopodite its tibiotarsus arises distad and diverges from the coaxial solenomere The Antillean species do not comprise a clade and are only distantly related rather than introductions they plausibly reflect ancestral occurrences on the ldquoproto-Antilleanrdquo terrain before it rifted from ldquoproto-South Americardquo in the CretaceousPaleocene with fragmentation isolating modern forms on their present islands Existing platyrhacid tribes are formally elevated to subfamilies as this category was omitted from recent taxonomies Without unequivocal evidence to the contrary geographically anomalous species should initially be regarded as indigenous rather than anthropochoric

Key words anthropochore Antilles Central America Guadeloupe Haiti Hispaniola Hoffmanorhacus Nannorrhacus New World Platyrhacidae Proaspis St Lucia South America West Indies

Introduction

The sudden passing of R L Hoffman in June 2012 deprived mankind of its most knowledgeable diplopodologist He was the only person who could take virtually any eugnathan milliped from any continent and put a tribal or lower name on it While he worked with nearly every eugnathan order he focused on the polydesmidan suborder Leptodesmidea (taxonomy according to Shelley [2003a] and Shear [2011]) with the largest-bodied ordinal representatives where he addressed five families comprehensively Chelodesmidae Gomphodesmidae Oxydesmidae Platyrhacidae and Xystodesmidae The base-level knowledge he established on the primarily Nearctic Xystodesmidae has been extensively enhanced and supplemented in the past 40 years and his voluminous treatises on the Afrotropical Gomphodesmidae and Oxydesmidae (Hoffman 1990 2005) are all-inclusive compendia He left the remaining familiesmdashthe Neo- and Afrotropical Chelodesmidae and the NeotropicalAustralasianOceanian Platyrhacidaemdashwith scores of tribal and generic treatments not having time to fulfill his desire to monograph both Perhaps over 95 of manrsquos knowledge of the last two assemblages died with him so diplopodologists must learn them anew and we offer this treatment of Antillean forms as a first step in advancing knowledge of Platyrhacidae without Dr Hoffmanrsquos encyclopedic insight He (Hoffman 1998) revised his previous

Accepted by W Shear 4 Jan 2013 published 15 Mar 2013 477

(Hoffman 1980) familial taxonomy (re)elevating Euryurinae and Aphelidesmini to familial ranks with Amplininae remaining a subfamily of the latter Platyrhacidae was left with seven tribes but no subfamilies so we elevate all present tribes (Hoffman 1998 2001a Jeekel 2007) these nine representing new statuses Barydesminae Diodontodesminae Erythrhacinae Hoplurorhachinae Phyodesminae Polydesmorhachinae Psammodesminae Psaphodesminae and Taphodesminae

The Antillean island arc of the eastern and northern Caribbean Sea harbors a curious and spotty mix of what we consider indigenous polydesmidans Of the four suborders (Shelley 2003a Shear 2011) Dalodesmidea is absent and Polydesmidea is widespread being known from all the Greater islands (Cuba Hispaniola Jamaica Puerto Rico) and all the Lessers that have been investigated plus the Bahamas (New Providence) which are actually in the Atlantic Ocean Antigua Dominica Grenada Grenadines (Carriacou) Guadeloupe (Basse-Terre) Marie Galante Tobago St Eustatius St Kitts St Lucia St Martin St Vincent and US Virgin Islands (St John) (Chamberlin 1918a Loomis 1933 1934 1936 1937 1938 1941a 1970 Veacutelez 1967a b de la Torre y Callejas 1974 Mauriegraves 1980 1988 Gonzalez Oliver and Golovatch 1990 Hoffman 1999 Peacuterez-Asso and Peacuterez-Gelabert 2001) Indigenous representatives of Leptodesmidea and Strongylosomatidea however occur sporadically particularly in the Lesser Antilles (Fig 1) Only one potentially native strongylosomatidean is known Mestosoma semirugosum (Pocock 1888) on Dominica (Pocock 1888 Chamberlin 1918a Hoffman 1960a 1999) and the last author suggested that it was introduced from an unknown source area Shelley and Golovatch (2011) however considered it indigenous and reflecting occurrence on the ldquoproto-Antilleanrdquo terrain before it rifted from ldquoproto-South Americardquo in the CretaceousPaleocene with isolation on present-day Dominica when the arc subsequently fragmented Antillean leptodesmideans comprise three families (Fig 1) Chelodesmidae in the Bahamas all the Greaters and Grenada and St Vincent Sphaeriodesmidae in Cuba Hispaniola (Dominican Republic Haiti) Jamaica and Guadeloupe (Basse-Terre) and Platyrhacidae in Hispaniola (Haiti) Guadeloupe (Basse-Terre) and St Lucia (Chamberlin 1918a Mauriegraves 1980 1988 Jeekel 1980 Hoffman 1980 1999 Shelley 2003b 2007 Peacuterez-Asso and Peacuterez-Gelabert 2001 Peacuterez-Gelabert 2008) As specified by Hoffman (1999) Hispaniolan sphaeriodesmids and platyrhacids are known only from Haiti and not from the adjacent and more heavily forested Dominican Republic Here we address Antillean platyrhacids which plausibly also arose on the ldquoproto-Antilleanrdquo terrain and became isolated on their present islands after rifting and fragmentation although Hoffman (1980 1999) and Jeekel (2007) thought that Nannorrhacus may represent a human introduction from the Indomalay Region Similar in their small body-sizes Antillean platyrhacids possess dissimilar gonopods and can only be interpreted as representing distinct monotypic genera

The first mention of a potential West-Indian platyrhacid was by Bollman (1888) who described Stenonia maculata from Cuba and commented that it ldquobelongs in the subgenus Stenonia (= Platyrhacus)rdquo Chamberlin (1918a) included ldquoPlatyrhacus () maculatusrdquo in his list of West Indian myriapods but Loomis (1950) and Hoffman (1999) referred the milliped to Pyrgodesmidae and the genus Docodesmus Cook 1896 Pocock (1894) described Platyrhacus luciae from St Lucia and Cook (1896) designated it type of the new genus Nannorrhacus Chamberlin (1918a) and Hoffman (1999) recognized P and N luciae respectively Hoffman (1960b) claimed the latter had been ldquofairly well describedrdquo but not in accordance with 21st century standards and Jeekel (1980) illustrated a gonopod and paranotum Loomis (1941b) proposed Proaspis aitia n gen n sp for three female platyrhacids in the southern peninsula of Haiti which was subsequently cited by Jeekel (1963) Hoffman (1999) Peacuterez-Asso and Peacuterez-Gelabert (2001) and Peacuterez-Gelabert (2008) Jeekel (1980) described and illustrated Proaspis sahlii from Basse-Terre Guadeloupe that Mauriegraves (1980) also illustrated while adding 25 new localities Though listing P sahlii Hoffman (1999) observed that congeneric status with P aitia could not be assumed until male topotypes of the latter became available and that acropodal differences between Jeekelrsquos and Mauriegravesrsquodrawings done from closely similar perspectives ldquostrongly suggest that two species are involvedrdquo

While perusing millipeds at the FSCA (acronyms below) in 2011 RMS discovered two platyrhacid samples one with five males collected in 1984 at Morne Formond a small mountain in Parc National de Macaya Sud Department Les Cayes Arrondissement near the western end of the southern Haitian peninsula The site is around 8ndash16 km (5ndash10 mi) SW of the type locality an unspecified site between Camp Perrin a municipality in the same arrondissement and department and Riviegravere Glace in GrandrsquoAnse Department We consider this sufficiently proximate to constitute near topotypes of Proaspis aitia particularly with an indefinite type locality and without other males Seventy-one years after its proposal we therefore establish the identity of P aitia and describe and illustrate males Since its biramous gonopodal telopodite is distinct from the uniramous one of P sahlii we propose

SHELLEY amp MARTINEZ-TORRES478 middot Zootaxa 3626 (4) copy 2013 Magnolia Press

Hoffmanorhacus n gen to accommodate the latter and redescribe N luciae to consolidate accounts of Antillean platyrhacids We conclude with biogeographical considerations based in part on new records (Appendix) suggest regions of potentially related continental forms and discuss characteristics of indigenous vs anthropochoric species Repository acronyms are BMNH The Natural History Museum London UK CAS California Academy of Sciences San Francisco USA FSCA Florida State Collection of Arthropods Gainesville USA NCBN Netherlands Centre for Biodiversity Naturalis Leiden LACM Los Angeles County Museum of Natural History Los Angeles California USA MPM Milwaukee Public Museum Milwaukee Wisconsin USA NCSM North Carolina State Museum of Natural Sciences Raleigh USA NMNH National Museum of Natural History Smithsonian Institution Washington DC USA UMSP Entomology Department University of Minnesota St Paul VMNH Virginia Museum of Natural History Martinsville USA

FIGURE 1 Indigenous occurrences of the polydesmidan suborders Leptodesmidea and Strongylosomatidea in the Antilles and Bahamas Chelodesmidae (Stars) Sphaeriodesmidae (Triangles) Platyrhacidae (Dots) Paradoxosomatidae (Diamond)

Taxonomy (taken in part from Hoffman 1980 1998 Shelley 2003a Shear 2011)

Order Polydesmida Pocock 1887

Suborder Leptodesmidea Broumllemann 1916

Superfamily Platyrhacoidea Pocock 1895

Family Platyrhacidae Pocock 1895

AntilleanWest Indian PlatyrhacidaeUndiscovered platyrhacids that may alter our perceptions and conclusions potentially occupy these andor

other Caribbean islands However we can only work with the known and the only detectable shared feature of the

Zootaxa 3626 (4) copy 2013 Magnolia Press middot 479PLATYRHACIDAE IN THE WEST INDIES

Antillean forms is relatively small body-size a nebulous and undefinable trait that clearly is not synapomorphic While the Antillean forms are significantly smaller than the large-bodied continental species of Barydesmus Nyssodesmus Psammodesmus and Tirodesmus all authored by Cook (1896) equally small-bodied forms occur in the continental range and particularly in Reserva Ntildeambi DMTrsquos research area in southwestern Colombia With different gonopodal configurations and tergal ornamentation varying from flat with polygonal areas in N luciae to heavily tuberculate in P aitia the Antillean platyrhacids are not united by synapomorphies and do not constitute a monophyletic group Hoffman (1980) placed Nannorrhacus in the nominate tribe now subfamily and cited Proaspis as ldquoPlatyrhacinae of uncertain status or tribal positionrdquo 32 years later we cannot assign either Proaspis or Hoffmanorhacus to a subfamily nor evaluate the position of Nannorrhacus Subfamilial assignments are thus deferred to DMTrsquos revision of New World platyrhacids as these catagories must be re-assessed in a collective revisionary context those of the Old World fauna need similar re-assessment

Epiproctal setal clusters In our low-power (lt200X) examinations we noted that the epiproctal setae in both P aitia and N luciae are actually clusters of 6ndash10 or so fine individual setae that we attempt to draw as such (Figs 4 14) the accurate term is thus ldquosetal clusterstuftsrdquo not ldquosetaerdquo or ldquosingle setaerdquo Checking through platyrhacid literature Chamberlin (1923 figs 115 123 129 154 194) sketched such clusters but did not mention them verbally Hoffman (1998133 fig 7) noted that ldquocompound setaerdquo occur on the labra legs and epiproct and considered them synapomorphic between Platyrhacidae and Aphelidesmidae which has priority over Amplininae for the familial name Shear (2008) examined epiprocts of representatives of 18 polydesmidan families and found spinnerets some with setal clusters on all of them Platyrhacidae was not one of these taxa but Silvestri (1903122 fig 202) drew the epiproct of Platyrhacus modigliani Silvestri 1895 revealing four linearly-arranged spinnerets on the caudal margin but not setal clusters The clusters we observed probably constitute evidence of platyrhacid spinnerets and in her further investigations DMT will investigate them under SEM

Terminology As gonopods are highly modified and specialized walking legs their terminology is applied to parts or regions of these secondary reproductive appendages Most particularly we employ ldquotelopoditerdquo referencing the part distal to the coxa and ldquoacropoditerdquo referencing that distal to the ldquoprefemurprefemoral regionrdquo the basal part of the telopodite possessing hairs and the pit in which the ldquoprostatic grooverdquo originates Thus the only difference is inclusion of ldquoprefemurrdquo in telopodite and its absence from acropodite

While some Old World platyrhacids for example taxa in Papua New Guinea and eastern Indonesia possess complex gonopodal telopodites with numerous distal branches (Hoffman 1997a b 2001b 2006) those of New World forms are comparatively simple as is their terminology All known forms are either uni- or biramous the latter without subdivision of any branch and with only a solenomere and a second terminal projection that may or may not be expanded or slightly modified both may be continuous or discontinuous with the axis of the telopodal stem By definition ldquosolenomererdquo is the branch or projection that carries the prostatic groove to its opening The name of the other branch is not set and we opt for the traditional term ldquotibiotarsusrdquo From published drawings (Attems 1999 pl 14 fig 336 1938 fig 229 Chamberlin 1923 pl 18 fig 118) and RMSrsquo examination of a male of Rhyphodesmus drurii (Gray 1832) from ldquoBarrikardquo (exact location unknown) Guyana (VMNH) its gonopods possess a secondary branch that arises basally on the acropodite is positioned ventral to the solenomere and parallels the latter a similar physical relationship to that of the ldquoendomererdquo in polydesmid (polydesmidean) gonopods like those of Scytonotus CL Koch 1847 and Calianotus Shelley 1997 (Shelley 1994 1997) Collecting or obtaining more males of Rhyphodesmus spp to fully assess gonopodal variation and the generic composition is an early priority of DMTrsquos future research

Key to species

1 Telopodite uniramous Basse-Terre Guadeloupe Hoffmanorhacus sahlii (Jeekel 1980)- Telopodite biramous possessing separate solenomere and tibiotarsus 22 Solenomere arising near telopodal midlength discontinuous from stem tibiotarsus long bent strongly laterad around mid-

length southern peninsula of Haiti Hispaniola Proaspis aitia Loomis 1941- Solenomere located distad continuous and coaxial with telopododal stem tibiotarsus short and sublinear diverging from

curved solenomere St Lucia Nannorrhacus luciae (Pocock 1894)

Genus Proaspis Loomis 1941

Proaspis Loomis 1941b193ndash194 Jeekel 196391ndash92 1971282 198085 (in key) Hoffman 195645 1960b19 (list) 1980164 1999399 Peacuterez-Asso and Peacuterez-Gelabert 200176 Peacuterez-Gelabert 200873

Type and only component species P aitia Loomis 1941b by original designationDiagnosis Heavily tuberculate Platyrhacidae paranota short flat and subparallel to substrate extending

essentially directly laterad lateral margins moderately incised ozopores and encircling rings submarginal half to full widths removed from lateral margins and just caudal to midwidths Gonopodal telopodite biramous extending through substantially more than one vertical plane without cingulum solenomere arising near midlength and extending dorsomediad discontinuous from stem tibiotarsus bending strongly and nearly directly laterad around 56 length narrowing and tapering at level of bend and also subterminally not overhanging solenomere

Distribution Known only from and possibly endemic to the western end of the southern peninsula of Haiti Hispaniola

Remarks Without males Hoffman (1980) could only place Proaspis under ldquoPlatyrhacinae of uncertain status or tribal positionrdquo With males we note that it differs from other New World taxa in the proximal position of the solenomere the narrow tibiotarsus and its strong lateral bend and the course of the prostatic groove which crosses from medial to lateral telopodal surfaces near midlengths These apparently strong apomorphies suggest separate subfamilial andor tribal statuses but such categories should not be erected on individual bases but in the context of the entire Western Hemisphere fauna In each occupied hemisphere all platyrhacid categories need to be re-evaluated collectively and simultaneously and more definitive diagnoses are in order DMT will assess those for American forms as part of a revision of the New World fauna and we therefore leave Proaspis without subfamilial assignment

Proaspis aitia Loomis 1941Figs 2ndash5

Proaspis aitia Loomis 1941b194 figs 10ndash12 Hoffman 1999399 Peacuterez-Asso and Peacuterez-Gelabert 200176 Jeekel 2007 91ndash92 Peacuterez-Gelabert 200873

Type specimen holotype (NMNH) now fragmented taken by HF Loomis 31 July 1940 along a roadside between Camp Perrin (Sud Department Les Cayes Arrondissement) and Riviegravere Glace (GrandrsquoAnse Department) Haiti Hispaniola Greater Antilles Two female paratypes were collected simultaneously (Loomis 1941b) but are now lost

Diagnosis With the characters of the genusColoration in preservative Topotypes randomly encrusted with soil Head tawny epicranium and genae

darker interantennal and frontal regions lighter Metaterga with broad light brown longitudinal stripe extending from collum to segment 18 paranota and epiproct darker brown

Male topotypes Least fragmented individual 337 mm long and 39 mm wide female 423 mm long Epicranium heavily tuberculate with small subequal mostly rounded tubercles terminating at level of antennae suture wide distinct Interantennal isthmus 11 mm wide tubercles much lower than on epicranium but with two higher conical ones demarcating frons Genae without impressions apices broadly rounded and extending beyond adjacent cranial margins more heavily tuberculate than relatively smooth frons width across genal apices 28 mm No evidence of facial setae other than clypeal (8-8) and labral (12-12) Antennae reaching back to caudal margin of 2nd tergite 1st article subglobose 2ndash6 clavate 7 short and truncate with 4 terminal sensory cones relative lengths of antennomeres 6gt5gt4gt3gt2gt1gt7 1ndash6 subequally pilose with variably long hairs 7 more densely hirsute with hairs partly obscuring terminal cones

Collum (Fig 2) 36 mm wide narrower than succeeding terga heavily tuberculate with row of larger ones caudal to anterior margin paranota margined by row of subequivalent tubercles with larger irregular ones on dorsal surface Proterga relatively smooth Metaterga of segments 2ndash18 (Figs 2ndash3) heavily endowed with irregular

tubercles some coalesced into clusters 19th metatergum narrower than preceding ones 33 mm wide Paranotal surfaces covered with dense sharp subconical tubercles 2nd paranotal margins lightly indented those of 3rd more

FIGURES 2ndash5 Proaspis aitia near topotypical male 2 left paranota and metaterga of collum and segments 2ndash3 dorsal view 3 left paranotum and metatergum of segment 10 dorsal view 4 epiproct dorsal view 5 telopodite of left gonopod lateral view s solenomere tt tibiotarsus

deeply and irregularly so 4th strongly incised and sawtoothed those of segments 5ndash18 variably and irregularly incised ozopores and surrounding rings (Fig 3) submarginal half to full widths removed from lateral margins and just caudal to midlines Epiproct (Fig 4) broad margins smooth dorsal surface lightly pustulate with 4 terminalsubterminal setal clusters arising from caudal margin or just back from same on dorsal surface

Pregonopodal sterna lightly pustulate without structural modifications Postgonopodal sterna becoming progressively more tuberculate caudad caudal margins concave Pregonopodal legs moderately hirsute more so distad gonapophyses short and broad arising from ventromedial surfaces of 2nd coxae Postgonopodal legs more strongly hirsute than pregonopodals becoming progressively more so caudad and on distal articles claws gently curved Paraproct surfaces moderately tuberculate with a large setiferous tubercle at 13 length and smaller one distad rims not elevated Hypoproct vaguely and irregularly semilunar surface lightly tuberculate caudal margin extending slightly in midline

Gonopodal aperture ca 14 mm wide and 07 mm long at midpoints located entirely in metazona margins strongly elevated above metazonal surface more so laterad anterior margin slightly indented in midline caudal margin with longer indentation Gonopods in situ small inconspicuous and fragile prefemora closely appressed to each other and nearly touching in midline telopodites diverging slightly and extending anteriad to level of anterior margin of 7th prozonum not overhanging 6th sternum solenomere apices overlapping in midline tibiotarsi bending strongly and nearly directly laterad Gonopod structure as follows (Fig 5) Coxae with around five lateral macrosetae arranged sublinearly Telopodite biramous extending through substantially more than one vertical plane prefemur comprising nearly half of length Solenomere arising near frac12 length just distal to prefemur linear for most of length and extending dorsomediad distally uncinate and apically subacuminate Tibiotarsus continuing linearly distal to origin of solenomere bending strongly abruptly and nearly directly laterad at frac12 length and not overhanging solenomere narrowing smoothly and continuously more so distad to acuminate tip Prostatic groove angling across medial surface basally crossing to lateral side at distal extremity of prefemur bending onto solenomere and opening apically

Variation The male near-topotypes agree closely with each otherEcology According to Loomis (1941b) the types were collected ldquobeneath loose grassrdquo No habitat notations

are provided for the near topotypical males but the female and juvenile were found on a ldquodead tree trunkrdquo Distribution Sud and probably also GrandrsquoAnse Departments in the western end of the southern peninsula of

Haiti Hispaniola Peacuterez-Asso and Peacuterez-Gelabert (2001) considered P aitia as endemic to Haiti Data for the near topotypes are as follows

Haiti Dept Sud Les Cayes Arrondissement Parc National de Macaya (18deg29rsquo04rdquo N 74deg08rsquo48rdquo W) Morne Formond South Slope Basin SSE Base Camp 2 1500ndash1530 m 5 and juv 1650 m SR Yocom (FSCA)

Remarks Proaspis aitia is shorter and narrower than its Antillean counterparts

Hoffmanorhacus new genus

Type and only component species Proaspis sahlii Jeekel 1980Diagnosis (adapted from Jeekel (1980)) Heavily tuberculate Platyrhacidae paranota relatively long flat and

subparallel to substrate extending essentially directly laterad from metazona lateral margins moderately scalloped ozopores and rings marginal and caudal to midwidths Gonopodal telopodite uniramous prefemur demarcated by distinct distal cingulum stem curving broadly anteriolaterad in substantially more than one vertical plane distal extremity curling dorsolaterad behind stem in medial view and looping back on itself to varying degrees depending on angle of view apically acuminate Distribution (Fig 11) Known only from and apparrently endemic to Basse-Terre Guadeloupe (Jeekel 1980 Mauriegraves 1980 1988)

Remarks Only so much variation and differentiation is possible in a uniramous telopodite without one or more additional branch(es) to change configuration position and angle of projection One species with uniramous gonopods is known to exist in the New World Amnestorhacus olonga (Silvestri 1899) (Hoffman and Martinez-Torres 2012) and the question arises as to whether sahlii is congeneric but after examining gonopods of two males of A ologona from Boyaca Dept Colombia (VMNH) we conclude that sahlii requires a new genus The two logically share ancestry and perhaps are sister-taxa but five anatomical differences summarized in table 1 seem significant enough to mandate different genera

TABLE 1 Comparison and contrast of Hoffmanorhacus and A ologona

The actual distal configuration and position varies in both taxa depending upon the angle of view whether the gonopod lies flat in a dish or the coxa or telopodite are tilted imperceptibly up or down which explains Hoffmanrsquos opinion (1999) that sahlii consisted of more than one species Even though the drawings of Jeekel (1980) and Mauriegraves (1980) were from medial aspects imperceptible tilting of the coxae or telopodites can convert one perspective of the distal extremity into another We also turned and manipulated the gonopods of the VMNH males of A olonga in every conceivable direction without achieving any of the distal configurations shown in figs 8ndash10 For all these reasons we are convinced that sahlii on Basse-Terre Guadeloupe is not congeneric with A olonga in central Colombia and we therefore erect Hoffmanorhacus to accommodate it Potential apomorphies for a clade containing these genera include the uniramous condition and the large broad telopodal prefemur that narrows and tapers distad

Hoffmanorhacus sahlii (Jeekel 1980) new combinationFigs 6ndash10

Proaspis sahlii Jeekel 198082ndash83 figs 1ndash3 200792 Mauriegraves 1988348 fig 1B Hoffman 1999399 Peacuterez-Asso and Peacuterez-Gelabert 200176 (text comments)

Proaspis () sahlii Mauriegraves 19801071ndash1072 figs 14ndash15

Type specimens (not examined) holotype allotype and 2 and 4 paratypes (NCBN) collected by F Sahli in January 1976 and September 1977 in Parc Naturel de Guadeloupe Basse-Terre Guadeloupe Lesser Antilles

Diagnosis With the characters of the genusDescription (Readers are referred to the detailed account in Jeekel [1980] we provide brief commentary on

somatic features and describe the gonopod to provide somewhat parallel accounts on all three Antillean platyrhacids)

Coxa with small field of macrosetae Telopodite uniramous without solenomere and tibiotarsus extending through substantially more than one vertical plane prefemur broad and stout basally narrowing distad and demarcated by distinct cingulum stem widening distal to cingulum then narrowing curving broadly dorsolaterad curling inward into arc looping back on itself and crossing dorsal to distalmost point of arc apically acuminate Prostatic groove running along medial side of prefemur crossing to lateral surface near cingulum and continuing to apical opening

Variation Not only does the distal configuration and curvature vary with imperceptible tiltings of the gonopod but so does the magnitude and visibility of the cingulum It is prominent in Fig 9 barely perceptible in fig 8 and imperceptible in fig 10

Ecology Apparently restricted to high altitude humid forests (Mauriegraves 1980) Distribution Known only from Basse-Terre Guadeloupe unknown from Grande-Terre Localities (Fig 11)

are as plotted by Mauriegraves (1988 fig 1B) Remarks Hoffmanorhacus sahlii is one of perhaps few diplopods with two patronyms in its official scientific

Hoffmanorhacus Amnestorhacus

Paranotal margins smooth to lightly scalloped Paranotal margins irregular to moderately jagged

Ozopores and surrounding rings located submarginally on paranota

Ozopores and rings located subcentrally on paranota

Telopodal prefemur demarcated distad by cingulum Prefemur not demarcated smoothly continuous with telopodal stem latter without cingulum

Telopodite curling smoothly and continuously distad Telopodite bending strongly and sharply distad

Distal telopodal extremity curling dorsolaterad behind stem in medial view

Distal telopodal extremity bending ventromediad in front of stem in medial view

FIGURES 6ndash10 Hoffmanorhacus sahlii 6ndash7 holotype reproduced with permission from Jeekel (1980) 6 left paranota and metaterga of collum and segments 2ndash3 dorsal view 7 left paranotum and metatergum of segment 10 dorsal view 8 right gonopod medial view 9ndash10 male from Matouba Basse-Terre Guadeloupe reproduced with permission from Mauriegraves (1980) 9 right gonopod medial view 10 the same anteriomedial view c cingulum

FIGURE 11 Occurrences of Hoffmanorhacus sahlii on Guadeloupe adapted and reproduced with permission from Mauriegraves (1988) BT Basse-Terre GT Grande-Terre

Jeekel (2007) suggested that H sahlii may be congeneric with Derodesmus haplopus Attems 1897 in Indonesia (Halmahera) and may hence be introduced from the Moluccan Islands We believe the differences are significant enough to mandate different genera for example though similar in being uniramous the telopodite of H sahlii is strong robust and heavily sclerotized whereas that of D haplopus is slender and fragile We therefore concur with Peacuterez-Asso and Peacuterez-Gelabert (2001) who considered it endemic to Basse-Terre and as explained in the generic account only one species inhabits the island

Genus Nannorrhacus Cook 1896

Nannorrhacus Cook 189652 Silvestri 1897649 (list) Jeekel 196388 1971274 Hoffman 195644 1960b18 (list) 1980163 1999398

Type and only component species Platyrhacus luciae Pocock 1894 by original designationDiagnosis Variably granular Platyrhacidae with at most only a few moderately high dorsal tubercles

paranota relatively long flat and subparallel to substrate extending essentially directly laterad lateral margins smooth and entire at most only lightly scalloped ozopores and rings submarginal half to full widths removed from lateral margins and just caudal to midwidths Gonopodal telopodite biramous extending through substantially

more than one vertical plane without cingulum distally bifurcate solenomere arising at 56 length (level of bifurcation) smoothly continuous with stem apically acuminate tibiotarsus angling generally dorsolaterad away from solenomere

Distribution Known only from two sites on St Lucia where it is probably endemic Its distribution on the island is unknown

Remarks Hoffman (1980) placed Nannorrhacus in the nominate tribe now subfamily but we remove it from this taxon and leave it unassigned

The double ldquorrdquo spelling of Nannorrhacus is correct because this was Pocockrsquos (1894) apparently intentional original spelling and there is no evidence of a lapsus necessitating emendation By contrast the double ldquorrdquo spelling of Platyrrhacus and Platyrrhacidae is incorrect because the original generic spelling by Koch (1847) was Platyrhacus with one ldquorrdquo

Attems (1899 1938) missed and apparently overlooked Nannorrhacus in both works he mentioned and characterized the species in the former but omitted even this from the latter

Nannorrhacus luciae (Pocock 1894)Figs 12ndash18

Platyrhacus luciae Pocock 1894 511 pl 39 figs 3 3andashd Chamberlin 1918a216Platyrrhacus luciae Attems 1899335Nannorrhacus luciae Loomis 193435 pl 1 fig 7 Hoffman 1999398 Peacuterez-Asso and Peacuterez-Gelabert 200176 (in text

comments) Jeekel 200791

Type specimen holotype (formerly BMNH but now lost) collected by G A Ramage on an unknown date prior to 1894 at Fond de Jacques St Lucia Lesser Antilles An extensive search in the BMNH holdings at RMSrsquo request failed to locate this specimen To the best of our knowledge the type locality and ldquoBar (Recte Barre) de lrsquoIsle above Castriesrdquo a hiking trail into the interior of the island that originates near Castries are the only definite records for N luciae and the distance along this trail is unknown Castries is around 18 km (12 mi) north of Fond de Jacques so from the standpoint of this small island only 435 km (272 mi) long the sites are not close together Given this distance and the uncertainty with the Barre de lrsquoIsle locality we do not designate a neotype to avoid potentially creating a nomenclatural problem in case more than one species inhabits the island St Lucia has not been sampled to the degree that Basse-Terre Guadeloupe has and while Nannorrhacus is probably monotypic we defer neotype designation to a time when it has been more thoroughly sampled additional localities are available and the exact number of components is known

Diagnosis With the characters of the genusColoration in preservative Randomly encrusted with soil Head creamish with small tubercles clypeus

darker Collum darker in midline metaterga generally cream-colored with irregular spots surrounded by gray proterga also cream-colored with small darker tubercles and circular brown spots in midlines

Male from St Lucia in general Ca 426 mm long and 64 mm wide Epicranium interantennal region frons and genae moderately granular suture deeply incised apically bifid isthmus 11 mm Genae relatively large with broad distinct medial impressions apices narrowly rounded width across 37 mm Facial setae absent except for clypeal (about 3-3) and labral (about 5-5) series these merging laterally and continuing for short distances along

genal margins Antennae extending back to caudal margin of 2nd tergite 1st article subglobose 2ndash5 clavate 6 cylindrical 7 short and truncate with 4 terminal sensory cones no other sensory structures evident relative lengths of antennomeres 6gt5=4=3=2gt1gt7 3ndash6 moderately pilose with variably long hairs 7 densely hirsute with shorter hairs

Collum (Fig 12) 41 mm wide surface densely granular with swollen elevated areas on each side of midline and row of low subflattened tubercles along anterior margin paranota essentially non-existent with caudolateral clusters of broadly rounded tubercles Proterga and metaterga of segments 2ndash16 (Figs 12ndash13) lightly granular with large flattened polygonal areas subdivided into smaller such areas larger ones becoming progressively more elevated caudad with smaller forming rows of low distinct tubercles along caudal margins of 18thndash19th metaterga Paranotal surfaces also with flattened dorsal polygonal areas becoming lightly tuberculate mediad on those of segments 18ndash19 margins smooth and nearly linear with fused barely perceptible scalloping ozopores and rings

positioned around frac12 widths from lateral margins and caudal to midwidths Epiproct (Fig 14) broad dorsal surface smooth with virtually no evidence of polygonal areas and two low tubercles giving rise to setal clusters margins also smooth with fused barely perceptible scalloping and posterior and lateral setal clusters

FIGURES 12ndash18 Nannorrhacus luciae 12 left paranota and metaterga of collum and segments 2ndash3 dorsal view 13 left paranotum and metatergum of segment 10 dorsal view 14 epiproct dorsal view 15 left gonopod medial view reproduced with permission from Jeekel (1980) 16 distal acropodal extremity of left gonopod of male from St Lucia in general lateral view 17 the same submedial view 18 the same anterior view s solenomere tt tibiotarsus

Sides of metazona lightly granular Pregonopodal sterna narrow surfaces smooth slightly elevated and lightly setose adjacent to coxae 6th sternum more elevated into knob-like structures and moderately setose adjacent to 6th

coxae Postgonopodal sterna wider and flattened surfaces smooth Pregonopodal legs moderately hirsute slightly

more so distad gonapophyses low inconspicuous elevations on 2nd coxae Postgonopodal legs slightly more hirsute and with longer hairs claws sublinear Paraproct surfaces smooth without polygonal areas with large tubercles at 13 and 23 lengths each giving rise to single macroseta rims not elevated Hypoproct large strongly trapezoidal extending for frac14 of paraproct lengths surface smooth and without polygonal areas caudal margin sublinear

Gonopodal aperture ca 11 mm wide and 07 mm long at midpoints located entirely in metazonum anterior margin flush with metazonal surface lateral margins elevating progressively caudad caudal margin slightly thickened and moderately elevated above metazonal surface Gonopods in situ strong and robust extending in subparallel arrangement over anterior margin of aperture and 7th prozonum tibiotarsal apices curving directly dorsad widely segregated from opposite member Gonopod structure (Figs 15ndash18) as follows Coxae with three

lateral macrosetae arranged sublinearly Telopodite biramous prefemur comprising about half of length curving broadly dorsad in essentially a single vertical plane around 23 length bifurcated at 56 length with solenomere continuing coaxially with stem and arc of loop curving inward and directed subventrad apically acuminate Tibiotarsus a separate branch arising dorsomediad at bifurcation diverging from solenomere and extending curvilinearly subdorsad narrowing sharply to acuminate tip Prostatic groove running along medial surface nearly to level of bifurcation crossing to lateral side and continuing along solenomere opening apically

Variation The tibiotarsus arises at approximately the same position in all males but its length relative to the solenomere and degree of curvature vary though to some degree this depends on the angle of view

Ecology UnknownDistribution Restricted to the island of St Lucia in the Lesser Antilles where to our knowledge Fond de

Jacques the type locality and ldquoBar (recte Barre) de lrsquoIsle above Castriesrdquo (Loomis 1934) are the only reported localities Jeekel (1980) did not provide a locality for the specimen(s) he illustrated The two known sites 18 km (12 mi) apart and in the southern and northern thirds of the island respectively suggest occurrence throughout St Lucia if the forms are conspecific which needs confirmation by male topotypes The following two samples were examined

St Lucia locality not specified February 1978 S Marshall (NCSM) and Barre de lrsquoIsle above Castries 2 February 1932 HF Loomis (FSCA)

Remarks Loomis (1934 pl 1 fig 7) published a dorsal-view photo of a maleJeekel (2007) believed the isolated occurrence of N luciae suggests introduced status but he acknowledged

that he could not associate it with an Indo-Australian platyrhacid From the gonopodal configuration he suggested affinity with the New World genus Barydesmus but we think this is distant at best (see relationships section) Furthermore the introduction would have had to take place prior to its discovery and probably no later than the 1880s This scenario seems unlikely and we believe N luciae is both indigenous and endemic to St Lucia Though its genus carries the ldquoNannohelliprdquo prefix meaning small N luciae is substantially larger-bodied than P aitia and its gonopods are also larger stouter and more strongly sclerotized

The Family Platyrhacidae in the New World

Distribution Hoffman (1978) stated that New World platyrhacids then platyrhacines ldquooccur only in the Andesrdquo He considered the Central American highlands an Andean extension and depicted the range extending from northern Costa Rica to western Peru which was incorrect when published Pocock (1894 1909) Broumllemann (1898) Cook (1896) Loomis (1941b) Chamberlin (1941 1950) Schubart (1947 1950) and Jeekel (1963) described or reported over 20 nominal species in several genera from Venezuela Guyana the Amazon lowlands of northeastern Peru and western and eastern Brazil in addition to St Lucia and Haiti Documentation from Auraacute Paraacute Est (Schubart 1947) established Platyrhacidae in the Amazon Delta of Brazil near the Atlantic Ocean implying occurrence to the north in French Guiana and Surinam that to our knowledge lack published records Apparently the only one from Venezuela is Platyrrhacus (sic) venezuelianus Broumllemann 1898 from ldquoLlanos du Venezuelardquo in general (Broumllemann 1898 Jeekel 2007) We know of no specific published localities in this country although RMS has seen samples from Neblina Massif on the border between Amazonas Ests Venezuela and Brazil The records of Platyrrhacus (sic) bilineatus (Lucas 1840) from ldquoAmazonesrdquo without further locality (Broumllemann 1909) ldquoPlatyrhacidae gen sprdquo from Auraacute Paraacute Est (Schubart 1947) and Platyrhacus (Tirodesmus) javarynus Schubart 1950 from along the Javari River Amazonas Est (Schubart 1950) seem the only authentic prior Brazilian localities Reports of ldquoPlatyrhacus scaber (Perty 1823)rdquo from Brazil in general and Minas Gerais Est (Broumllemann 1909) and P rufipes Koch 1847 from ldquoUnggewiss wahrscheinlich Brazilienrdquo that Koch (1863) and Broumllemann (1909) reiterated are problematic and for complex nomenclatural reasons have been discounted by Hoffman (1956 1974 1980) and Jeekel (1963 1971) The only definite and authentic Brazilian records that we can find are those of Schubart (1947 1950) from Amazonas and Paraacute Ests in the west and east respectively

We project a New World familial distribution (Fig 19) that includes the Antillean species and incorporates new localities from Brazil Colombia and Peru (Appendix) the dashed line reflects the unknown Brazilian range south of the Amazon We believe Platyrhacidae occupies a subcontinuous range on the continental land mass that encloses roughly the northern frac14 of South America and extends northward through southern Central America to

southeastern Nicaragua encompassing all or essentially all of Costa Rica Panama Colombia Ecuador Venezuela Guyana Surinam and French Guiana The northernmost locality is an unspecified site on Rio Machuca Rio San Juan Dept Nicaragua (Chamberlin 1922 Loomis 1968 Hoffman 1999) the easternmost is Auraacute Paraacute Est Brazil (Schubart 1947) and the southernmost appears to be Pampa Hermosa Junin Dept Peru (Chamberlin 1941) Platyrhacidae surely ranges farther south in Peru as well as Brazil and may even spread into northern Bolivia but we doubt it extends as far south as the southcentral locality of the order Glomeridesmida otherwise sympatric in South America (Shelley and Golovatch 2011 Shelley 2011) The family inhabits only southernmost Central America and is thus congruent with Epinannolenidea (Spirostreptida) and Paradoxosomatidae (Polydesmida Strongylosomatidea) (Loomis 1972 Hoffman 1977 1999 Shelley and Golovatch 2011) The last authors documented two distribution patterns for Gondwanan taxa in Central America a short one encompassing only Panama and part of Costa Rica shown by the aforementioned taxa and a long one extending to at least southern Mexico demonstrated by Glomeridesmida Siphonophorida Spirobolida Spirostreptidea (Spirostreptida) and Stemmiulida adjusted northward in Mexico by Shelley et al (2012) These distinct patterns with 1600 km (1000 mi) separating the northernmost locality of the first and the southernmost northern record of the latter indicate separate penetrations of ldquoproto-NorthCentral Americardquo Ancestors exhibiting the longer pattern probably entered around 306 million years ago (mya) in the Late Carboniferous Period Paleozoic Era when ldquoEuramericardquo merged with the ldquoproto-South Americardquo part of the existing Gondwana while those exhibiting the shorter entered 301 my later when the Panamanian Portal closed in the Pliocene Period Coenozoic Era 5 mya Consequently Epinannolenidea and Paradoxosomatidae have only had time to disperse through present-day Panama and half of Costa Rica Platyrhacidaersquos distribution conforms to this pattern and the ~144 km (90 mi) farther northward that it has spread logically reflects the greater motility of these large-bodied polydesmidans Platyrhacidae thus constitutes a Gondwanan taxon whose occurrence in Central America represents dispersal from a South American source area dating back 5 my to closure of the isthmus of Panama and completion of the land bridge between South and North America

Relationships The origins and affinities of the Antillean platyrhacids have baffled diplopodologists for decades as evidenced by at least four un- or minimally substantiated statements that one or more must represent introductions from the Old WorldIndomalay fauna (Jeekel 1963 2007 Hoffman 1980 1999) It is easy to dismiss or rationalize geographically anomalous occurrences as anthropochores from distant parts of the world but analysis and serious reflection are needed For example Hoffman (1999) suggested that the occurrence of Amphelictogon subterraneus bahamiensis Chamberlin 1918 on Andros Island Bahamas represented such but with increasing discoveries of Bahamian chelodesmids Shelley (2003b 2007) described how they plausibly reflect natural range expansion and indigenous occurrence ndash rafting once from Cuba during the Pleistocene reproducing and dispersing through the single ldquoGrand Bankrdquo island and becoming isolated on todayrsquos islands (Andros Cat Eleuthera and New Providence) as sea levels rose in the Post-Pleistocene Period The limited occurrences of Bahamian chelodesmids and Antillean leptodesmideans are inconsistent with the population explosions that truly anthropochoric species often undergo they also disseminate rapidly through human commerce quickly overrun small to moderate-size islands like the Lesser Antilles and even become household pests Unquestionably exogenous millipeds like Rhinotus purpureus (Pocock 1894) (Polyzoniida Siphonotidae) Trigoniulus corallinus(Gervais 1847) (Spirobolida Trigoniulidae) and Asiomorpha coarctata (Saussure 1860) (Polydesmida Paradoxosomatidae) are widespread to virtually ubiquitous on Caribbean Islands (Loomis 1934 1936 Jeekel 1963 Hoffman et al 1996 2002 Hoffman 1999 Shelley and Lehtinen 1999) and no evidence exists that leptodesmideans with limited distributions have achieved the notoriety of true introductions We do not know definitely whether N luciae is widespread or restricted on St Lucia but it appears to be endemic likewise there is no evidence that P aitia and H sahlii have spread beyond their documented ranges on Hispaniola and Guadeloupe Indeed Mauriegraves (1980 1988) stated that the latter occurs only in ldquola forecirct humide de Basse-Terrerdquo while a true introduction would certainly inhabit closely proximate Grande-Terre Likewise P aitia is not known elsewhere in Haiti or in the adjacent and more heavily forested Dominican Republic (Hoffman 1999 Peacuterez-Asso and Peacuterez-Gelabert 2001 Peacuterez-Gelabert 2008) All three platyrhacids may await discovery elsewhere but we think sufficient sampling has taken place that at least one additional occurrence of one species would exist by now if they were truly introduced That this has not happened indicates that hypotheses of true endemism must be formulated and their substantially different gonopods indicate that affinities must be sought elsewhere in the Western Hemisphere itself Phylogenetic analyses of the Antillean forms alone is impossible such must occur in the analytical context

of the total New World fauna which requires substantially more material from parts of South America and is beyond our present scope DMT will conduct these as part of her revision but in the meantime no harm is done by investigating whether geographical considerations provide clues that enable some degree of understanding

FIGURE 19 Known and projected distribution of Platyrhacidae in the Western Hemisphere Diamond Proaspis aitia Star Hoffmanorhacus sahlii Triangle Nannorrhacus luciae Dots new and borderline records in the continental range The dashed line denotes the uncertain southern range boundary through the heart of Brazil

The sporadic Antillean platyrhacid occurrences are matched in the Lesser Antilles by the equally anomalous ones of Paradoxosomatidae on Dominica Chelodesmidae on Grenada and St Vincent and Sphaeriodesmidae on Guadeloupe (Pocock 1888 Hoffman 1960a 1999 Mauriegraves 1980) (Fig 1) Only the first has been rationalized as an introduction (Hoffman 1960a 1999) but Shelley and Golovatch (2011) theorized that its ancestor inhabited the central ldquoproto-Antilleanrdquo region of Gondwana before it rifted from ldquoproto-South Americardquo in the Cretaceous Paleocene ~66 mya drifted northward rotated and became isolated on present-day Dominica during subsequent fragmentation We believe this scenario plausibly explains the origins of Antillean platyrhacids and since N luciaeis the southernmost its source area was plausibly the easternsouthernmost prior to rifting (Fig 20) Likewise that of H sahlii occupying a central position today was plausibly central prior to rifting and that of the westernmost P aitia was probably the westernmost (Fig 20) consequently affinities may lie in these areas of the present continental range Starting with N luciae with its continuous coaxial solenomere and distolateral tibiotarsus the only known continental species whose anatomical features are plausibly synapomorphic is Rhyphodesmus drurii(Gray 1832) in Guyana (Jeekel 1963 2007) Its smoothly curving solenomere forms the telopodal stem and there

is a separate shorter endomere-like ldquotibiotarsusrdquo arising proximad and curving parallel to the latter (Attems 199982ndash83 pl 14 fig 336 1938213 fig 229 Chamberlin 1923 44ndash45 pl 18 fig 118) Though substantially different this platyrhacid is closest anatomically to N luciae among forms known today and it is also the easternmost and the most geographically proximate to the position where ldquoproto-St Luciardquo was attached prior to rifting Substantial divergence has surely taken place in both forms but N luciae evolving in isolation has likely diverged farther from the ancestral condition and the position of its tibiotarsus seems to represent distal relocation along the telopodal stem The telopodal condition of Nannorrhacus has more in common with Rhyphodesmus than with more western continental taxa like Barydesmus Psammodesmus and Nyssodesmus and the two plausibly arose from a shared ancestor in the area of present-day Guyana (under-sampled) and SurinamFrench Guiana (wholly uninvestigated) (Fig 20) Sampling in the three northeasternmost South American countries will reveal whether forms closer to the condition in Nannorrhacus occupy this area today

FIGURE 20 Plausible areas of Antillean platyrhacid affinities with forms on the continental land mass Diamond Proaspis aitia Star Hoffmanorhacus sahlii Triangle Nannorrhacus luciae Dots general area of occurrence of Arcidesmus ologona in central Colombia

Hoffmanorhacus is readily distinguished from the other Antillean taxa by its uniramous gonopods Its telopodite is smoothly continuous without stub-like vestiges that could signify a lost branch so if the uniramous condition is derived it has existed long enough that traces of its origin have vanished and hence it probably antedates rifting of the Antillean arc One platyrhacid with uniramous telopodites logically deriving from the dichotomy yielding ancestral Hoffmanorhacus inhabits South America Amnestorhacus olonga in the Colombian interior It is removed from the coastal region that adjoined this part of the Antillean terrain before rifting but lies directly in line with Guadeloupe and the central Venezuelan Coast (Fig 20) where future sampling should focus (Hoffman and Martinez-Torres 2012)

Around the Antillean Arc Haiti is farther from Guadeloupe (~1491 km (932 mi)) than the latter is from St Lucia (~256 km (160 mi)) Consequently the affinities of Proaspis logically lie farther westward in South America but the dichotomy that led to ancestral Proaspis had to take place in ldquoproto-South Americardquo not Central America because the rifting occurred some 60 my before the Panamanian portal closed Though sharing the biramous condition with Barydesmus Psammodesmus and Nyssodesmus Proaspis is unique in that its tibiotarsus

is fragile slender and bends abruptly laterad and the solenomere arises laterally near telopodal midlength Accordingly the prostatic groove crosses from medial to lateral sides of the stem around 13 length rather than running entirely along the former as in Psammodesmus With notable differences the features of Proaspis resemble those of Nyssodesmus and Barydesmus (Table 2) we believe they likely share ancestry and that the source area of ancestral Proaspis plausibly lies in northwestern Colombia between the Caribbean Sea and the juncture with Panama and Central America (Fig 20)

TABLE 2 Acropodal differences between Proaspis Barydesmus and Nyssodesmus

Conclusion

Our independent and collaborative studies on New and Old World platyrhacids are only beginning but we can state even now that it is a complex monophyletic taxon that may be replete with homoplasies and convergences preliminary studies suggest that the uni- or biramous telopodal conditions may have evolved independently in each hemisphere As mentioned our distinguished predecessors RL Hoffman and CAW Jeekel repeatedly speculated that Antillean platyrhacids may be anthropochores and even suggested Old World source areas but all of these statements are baseless and lack scientific merit To our dismay we discovered during a final reference check that Jeekel (2007) expanded his prior comments (Jeekel 1963) and with no supportive evidence speculated that Rhyphodesmus drurii in South America may also be such and even identified a probable source area ldquoIt does not seem impossible that Rhyphodesmus drurii was introduced into Guyana from southern Burma or the Malayan Peninsulardquo He thought discovery in the latter areas was possible but then hedged and admitted that ldquothe recorded Guyana localities do not all seem to be typical synanthrope sitesrdquo It is one thing to make such statements for species like N luciae on small to minute islands that are typically overrun by anthropochores but quite another to do so for ones on continental land masses where indigenous forms overwhelmingly dominate We therefore emphasize that no credible scientific basis exists to think that any platyrhacid in either hemisphere was introduced by man from the other hemisphere Logically the original importations would have had to occur no later than the mid-1800s to allow time for population sizes to develop Columbus did not bring them to the West Indies because platyrhacids do not occur in Europe and we doubt that clipper ships were sailing around Cape Horn inadvertently carrying them but no other millipeds between the two familial range areas These statements seem to derive from the discovery of unexpected forms in unanticipated areas Without obvious explanations as to origins and affinities it is easy to rationalize them as ldquointroductionsrdquo and Hoffman and Jeekel did so

We submit that without overt evidence of introduced status (synanthropic habits large population sizes widespread or ubiquitous distributions) species that appear geographically illogical rarely if ever are anthropochoric The ldquosurpriserdquo that registers in our minds actually reflects our lack of comprehension of the taxon Instead of dismissing inexplicably anomalous species as introductions study and investigation are needed to gain the knowledge that will illuminate them We further submit that upon discovery of a ldquosurpriserdquo form and barring obvious attributes of an introduction the initial assumption should always be indigenous occurrence In the United

Proaspis Loomis 1941 Barydesmus Cook 1896 Nyssodesmus Cook 1896

Tibiotarsus relatively long and narrow constituting frac12 or more of acropodal length of subequal breadth throughout except for distal tapering

Tibiotarsus relatively short and moderately broad constituting no more than frac14 of acropodal length variably but only moderately expanded

Tibiotarsus short to moderately long varying from an immeasurably short tip to perhaps 13 of acropodal length moderately to broadly expanded

Tibiotarsus bent strongly and abruptly sublaterad near midlength

Distal extremity of acropodite upright or curving smoothly and variably subdorsad

Distal extremity of acropodite bent or curving strongly and abruptly sublaterad

Solenomere long slender and apically curled directed submediad and arising near midlength of lateral acropodal surface

Solenomere variably short and spiniform directed anteriodorsad and arising distad from dorsal surface of acropodal stem

Solenomere variably long and slender directed laterad and arising distally or subapically from distodorsal surface of acropodal stem bending under and obscured by distal acropodal expansion or clearly separated from latter

States for example man has recently and unintentionally introduced the southern Appalachian millipeds Cleidogona nantahala Shear 1972 (Chordeumatida Cleidogonidae) and Boraria stricta (Broumllemann 1896) and B infesta (Chamberlin 1918) (Polydesmida Xystodesmidae) into New England (Shear and Shelley 2004 Shelley et al 2011) The first and last occur exclusively in synanthropic situations and B stricta has exploded in Mianus River Gorge Westchester County New York close to upscale residential areas With overwhelming evidence of anthropochoric status nobody questions whether the New England populations may be indigenous Similarly the enigmatic genus Choctella Chamberlin 1918 (Spirostreptida Choctellidae) is a geographical anomaly in the Cumberland Plateau of Tennessee and Alabama (Chamberlin 1918b Chamberlin and Hoffman 1950 1958 Hoffman 1965 1999 Shelley and Golovatch 2011) but nobody has dismissed it as an introduction from a distant part of the planet Unquestionably indigenous its species have not undergone population blooms occur in both urban and natural habitats and have not spread beyond their documented areas However the similar geographical anomalies of a chelodesmid in the Bahamas and platyrhacids in Haiti and St Lucia inexplicably caused our deceased colleagues to speculate that they were introduced and Jeekel (2007) even suggested such for South American platyrhacids that occur in remote jungle well beyond significant human influence and exhibit variation consistent with natural occurrence We are convinced that the West Indian platyrhacids and the continental forms of Rhyphodesmus are indigenous and probably endemic to their respective areas and suggest herein an explanatory scenario to be confirmed refuted expanded or modified in future investigations As with Bahamian chelodesmids a presently unknown explanation must exist but research is necessary to elucidate it and formulate scientifically credible hypotheses New milliped introductions will continually arise through increasing human commerce but the temptation to dismiss every geographically anomalous species as anthropochoric must be resisted In the absence of convincing evidence to this effect the initial assumption should always be indigenous occurrence

Acknowledgments

We thank SI Golovatch and WA Shear for presubmission and editorial reviews and GB Edwards (FSCA) for access to and loan of the near-topotypical samples of P aitia and other platyrhacids in this collection Dana DeRoche (NMNH) and BJH Brugge (NCBN) provided data on the types of P aitia and P sahlii respectively and J Beccaloni searched in vain for the type of Platyrhacus luciae in the BMNH Access to and loans of other platyrhacid material were courtesy of the following curators and collection managers CE Griswold (CAS) the late CL Hogue (LACM) JP Jass (MPM) JA Coddington (NMNH) R Hozenthal and PK Clausen (UMSP) and H Cartmell (VMNH) Permissions to reproduce published figures and maps of H sahlii and N luciae were granted by J-P Mauriegraves and H de Jong and E Floacuterez Daza allowed access to the ICN database Finally we must acknowledge the multitude of detailed anatomical works on platyrhacids by RL Hoffman the Grand Master of Diplopodology and 41 years of wonderfully insightful and illuminating discussions with RMS that manifestly enhanced his grasp of this great arthropod class As with most modern diplopod studies the present work is possible only because of the multitude of tangible and intangible contributions by Dr Hoffman on both platyrhacids and diplopods in general

Literature cited

Attems C (1899) System der polydesmiden II Theil Denkschriften der Mathematisch-Naturwissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschaften Wien 68 251ndash436

Attems C (1938) Myriapoda 3 Polydesmoidea II Fam Leptodesmidae Platyrhacidae Oxydesmidae Gomphodesmidae Das Tierreich 69 1ndash487

Bollman CH (1888) Notes on a collection of Myriapoda from Cuba Proceedings of the United States National Museum 11 335ndash338 httpdxdoiorg105479si0096380111-720335

Broumllemann HW (1898) Myriapodes du haut et bas Sarare (Venezuela) donneacutes par M F Geay au Museacuteum drsquoHistoire Naturelle de Paris Annales de la Socieacuteteacute Entomologique de France 68 314ndash334

Broumllemann HW (1909) Os myriapodos do Brazil Catalogos da fauna Brazileira editados pelo Museu Paulista S Paulo ndash Brazil 2 1ndash94 + indashxx

Causey NB (1965) Additions to the milliped genus Siphonoconus (Siphonotidae Polyzoniida) Proceedings of the Louisiana

Academy of Sciences 28 51ndash55Chamberlin RV (1918a) The Chilopoda and Diplopoda of the West Indies Bulletin of the Museum of Comparative Zoology at

Harvard College 62(5) 151ndash262 Chamberlin RV (1918b) Myriapods from Nashville Tennessee Psyche 25 23ndash30

httpdxdoiorg101155191865230Chamberlin RV (1922) The millipeds of Central America Proceedings of the United States National Museum 60(8) 1ndash75

httpdxdoiorg105479si0096380160-24031Chamberlin RV (1923) Results of the Bryant Walker Expeditions of the University of Michigan to Colombia 1913 and

British Guiana 1914 Occasional Papers of the Museum of Zoology University of Michigan 133 1ndash143Chamberlin RV (1941) On a collection of millipedes and centipedes from northeastern Peru Bulletin of the American

Museum of Natural History 78(7) 473ndash535Chamberlin RV (1950) Neotropical chilopods and diplopods in the collections of the Department of Tropical Research New

York Zoological Society Zoologica 35(2) 133ndash144 Chamberlin RV and RL Hoffman (1950) Some families and genera of North American diplopods Chicago Academy of

Science Natural History Miscellanea 71 1ndash7Chamberlin RV and RL Hoffman 1958 Checklist of the millipeds of North America United States National Museum

Bulletin 212 1ndash236 httpdxdoiorg105479si03629236212Cook OF (1896) New American Platyrrhacidae Brandtia 12 51ndash54de la Torre Y Callejas SL (1974) Lista preliminar de los diplopodos (Miriaacutepoda Diplopoda) de Cuba Ciencias Biologicas

42(6) 1ndash16Gonzalez Oliver R and SI Golovatch (1990) Cataacutelogo de los Diploacutepodos de Cuba Editorial Academia La Habana 37 ppHoffman RL (1956) Generic names in the family Platyrhacidae and their type species with a consideration of the status of

Stenonia Gray 1842 Proceedings of the Biological Society of Washington 69 41ndash52Hoffman RL (1960a) Millipeds from Dominica British West Indies Proceedings of the United States National Museum 111

33ndash41Hoffman RL (1960b) A fourth contribution to the knowledge of Neotropical platyrhacid millipeds (Diplopoda Polydesmida)

Proceedings of the United States National Museum 111 17ndash31Hoffman RL (1965) A second species in the diplopod genus Choctella (Spirostreptida Choctellidae) Proceedings of the

Biological Society of Washington 78 55ndash58Hoffman RL (1974) Platyrhacus Koch 1847 (Diplopoda) Petition for the designation by the International Commission of a

type-species in harmony with accustomed usage ZN(S) 2078 Bulletin of Zoological Nomenclature 31(4) 249ndash251Hoffman RL (1977) The milliped genus Mestosoma in Costa Rica (Polydesmida Paradoxosomatidae) Studies on

Neotropical Fauna and Environment 12 207ndash215 httpdxdoiorg10108001650527709360520

Hoffman RL (1978) On the classification and phylogeny of chelodesmoid Diplopoda Abhandlungen und Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg Neue Folge 2122 21ndash31

Hoffman RL (1980 [1979]) Classification of the Diplopoda Museacuteum dHistoire Naturelle Genegraveve Suisse 237 ppHoffman RL (1990) Myriapoda 4 Polydesmida Oxydesmidae Das Tierreich Teil 107 Walter de Gruyter Berlin 512 ppHoffman RL (1997a) A new psaphodesmine genus from New Guinea (Polydesmida Platyrhacidae) Myriapodologica 4(12)

101ndash106Hoffman RL (1997b) Synopsis of the milliped genus Parazodesmus (Polydesmida Platyrhacidae) Steenstrupia 23 21ndash36Hoffman RL (1998) Reassessment of the Platyrhacidae a family of polydesmidan millipeds Myriapodologica 5 125ndash141Hoffman RL (1999) Checklist of the millipeds of North and Middle America Virginia Museum of Natural History Special

Publication 8 1ndash584Hoffman RL (2001a) A synopsis of the platyrhacid millipeds of Borneo (Diplopoda Polydesmida Platyrhacidae) Revue

Suisse de Zoologie 108 403ndash440Hoffman RL (2001b) Synopsis of Zodesmus a genus of Papuan millipeds (Polydesmida Platyrhacidae Psaphodesmini)

Myriapodologica 7(8) 59ndash76Hoffman RL (2005) Monograph of the Gomphodesmidae a family of African polydesmoid millipeds Naturhistorisches

Museum Wien 537 ppHoffman RL (2006) Four new psaphodesmine genera from the Papuan Region (Polydesmida Platyrhacidae)

Myriapodologica 8(10) 93ndash107Hoffman RL and D Martinez-Torres (2013) Amnestorhacus a new genus in the Neotropical diplopod fauna based on the

forgotten species Arcidesmus ologona Silvestri 1898 (Polydesmida Platyrhacidae) Annali del Museo Civico di Storia Naturale ldquoGiacomo Doriardquo 104 131ndash140

Hoffman RL SI Golovatch J Adis and JW de Morais (1996) Practical keys to the orders and families of millipedes of the Neotropical region (Myriapoda Diplopoda) Amazoniana 14(12) 1ndash35

Hoffman RL SI Golovatch J Adis and JW de Morais (2002) Diplopoda pp 505ndash533 In Adis J (ed) Amazonian Arachnida and Myriapoda Pensoft Publishers Sofia Bulgaria 590 pp

Jeekel CAW (1963) Diplopoda of Guiana (1ndash5) Studies Fauna Surinam 4 1ndash157 Jeekel CAW (1971) Nomenclator generum et familiarum Diplopodorum A list of the genus and family-group names in the

class Diplopoda from the 10th edition of Linnaeus 1758 to the end of 1957 Monografieeumln van de Nederlandse Entomologische Vereniging 5 1ndash412 + indashxii

Jeekel CAW (1980) A new species of the family Platyrhacidae from Guadeloupe West Indies (Diplopoda Polydesmida) Bulletin Zoologisch Museum Universiteit van Amsterdam 7(9) 81ndash87

Jeekel CAW (2007) An annotated bibliographical catalogue of the Indo-Australian Platyrhacidae (Diplopoda Polydesmida) Myriapod Memoranda 10 49ndash101

Koch CL (1847) System der Myriapoden pp 131ndash132 In Panzer and Herrich-Schaumlffer Kritische Revision der Insectenfauna Deutschlands 3 1ndash1496

Koch CL (1863) Die Myriapoden Getreu nach der Naturabgebildet und Beschrieben Erster Band Halle Regensburg 134 pp httpdxdoiorg105962bhltitle10639

Loomis HF (1933) Three new Cuban millipeds with notes on two little-known species Bulletin of the Museum of Comparative Zoology at Harvard College 75(9) 357ndash363

Loomis HF (1934) Millipeds of the West Indies and Guiana collected by the Allison V Armour expedition in 1932 Smithsonian Miscellaneous Collections 89(14) 1ndash69

Loomis HF (1936) The millipeds of Hispaniola with descriptions of a new family new genera and new species Bulletin of the Museum of Comparative Zoology at Harvard College 80(1) 3ndash191

Loomis HF (1937) New Jamaican and Cuban millipeds with notes on several other species Bulletin of the Museum of Comparative Zoology at Harvard College 80(3) 215ndash228

Loomis HF (1938) New and noteworthy millipeds from Cuba collected by Dr PJ Darlington in 1936 Bulletin of the Museum of Comparative Zoology at Harvard College 83(6) 427ndash480

Loomis HF (1941a) A new Cuban milliped with notes and drawings of other West Indian species Psyche 48 35ndash39 httpdxdoiorg101155194156917

Loomis HF (1941b) New genera and species of millipeds from the southern peninsula of Haiti Journal of the Washington Academy of Science 31 188ndash195

Loomis HF (1950) Synonymy of some native American and introduced millipeds Journal of the Washington Academy of Science 40 164ndash166

Loomis HF (1968) A checklist of the millipeds of Mexico and Central America United States National Museum Bulletin 266 1ndash137 httpdxdoiorg105479si03629236266

Loomis HF (1970) Millipeds of St John U S Virgin Islands and a new species from Puerto Rico Florida Entomologist 53(3) 129ndash134 httpdxdoiorg1023073493357

Loomis HF (1972) Millipeds from the Atlantic lowlands of Costa Rica Florida Entomologist 55(3) 185ndash206Mauriegraves J-P (1980) Diplopodes chilognathes de la Guadeloupe et ses deacutependances Bulletin du Museacuteum National drsquoHistoire

Naturelle 4e seacuterie section A 2(4) 1059ndash1111Mauriegraves J-P (1988) Les modes de peuplement des iles oceaniques par les diplopodes Le cas des Antilles Franccedilaises Bulletin

de la Socieacuteteacute Zoologique de France 112(3ndash4) 343ndash353Peacuterez-Asso AR amp Peacuterez-Gelabert DE (2001) Checklist of the millipeds (Diplopoda) of Hispaniola Boletin de la Sociedad

Entomologica Aragonesa 28 67ndash80Peacuterez-Gelabert DE (2008) Arthropods of Hispaniola (Dominican Republic and Haiti) A checklist and bibliography Zootaxa

1831 1ndash530Pocock RI (1888) Contributions to our knowledge of the Myriopoda of Dominica Annals and Magazine of Natural History

Series 6 2 472ndash483Pocock RI (1894) Contributions to our knowledge of the arthropod fauna of the West Indies Part III Diplopoda and

Malacopoda with a supplement on the Arachnida of the class Pedipalpi Journal of the Linnean Society (Zoology) 24 473ndash544 httpdxdoiorg101111j1096-36421894tb02494x

Pocock RI (1909) Diplopoda Biologia Centrali-Americana 137ndash147 Schubart O (1947) Os Diplopoda da viagem do naturalista antenor Leitatildeo de Carvalho aos rios Araguaia e Amazonas em 1939

e 1940 Boletim do Museu Nacionalo Nova Seacuterie Zoologia 82 1ndash74Schubart O (1950) Novos diploacutepodos do Brasil Papeacuteis Avulsos do Departamento de Zoologia 9(11) 145ndash157 Shear WA (2008) Spinnerets in the milliped order Polydesmida and the phylogenetic significance of spinnerets in millipeds

(Diplopoda) International Journal of Myriapodology 2 123ndash146 httpdxdoiorg101163187525408X395904

Shear WA (2011) Class Diplopoda de Blainville in Gervais 1844 pp 159ndash164 In Zhang Z-Q (Ed) Animal biodiversity An outline of higher-level classification and survey of taxonomic richness Zootaxa 3148 1ndash237

Shear WA amp Shelley RM (2004) Introduction of the milliped Cleidogona nantahala Shear 1972 in New England (Chordeumatida Cleidogonidae) Entomological News 115(2) 88ndash94

Shelley RM (1994 [1993]) Revision of the milliped genus Scytonotus Koch (Polydesmida Polydesmidae) Brimleyana 19 1ndash60

Shelley RM (1997) The identity of Polydesmus sastianus Chamberlin proposal of a new milliped genus and remarks on the

identity of Phreatodesmus hastingsus (Chamberlin) (Polydesmida Polydesmidae) Myriapodologica 4(8) 59ndash67Shelley RM (2003a [2002]) A revised annotated family-level classification of the Diplopoda Arthropoda Selecta 11(3)

187ndash207Shelley RM (2003b) Redescription of the milliped Amphelictogon subterraneus bahamiensis Chamberlin 1918 with an

assessment of the family Chelodesmidae in the Bahamas (Polydesmida Leptodesmidea) Zootaxa 180 1ndash8Shelley RM (2007) A fourth Bahamian record of the milliped Amphelictogon subterraneus bahamiensis Chamberlin 1918

(Polydesmida Chelodesmidae) Entomological News 118(3) 323ndash324 httpdxdoiorg1031570013-872X(2007)118[323AFBROT]20CO2

Shelley RM (2011) The milliped order Glomeridesmida (Diplopoda Pentazonia Limacomorpha) in Oceania the East Indies and southeastern Asia first records from Palau the Philippines Vanuatu New Britain the Island of New Guinea Cambodia Thailand and Borneo and Sulawesi Indonesia Insecta Mundi 0196 1ndash11

Shelley RM and SI Golovatch (2011) Atlas of myriapod biogeography I Indigenous ordinal and supra-ordinal distributions in the Diplopoda Perspectives on taxon origins and ages and a hypothesis on the origin and early evolution of the class Insecta Mundi 0158 1ndash134

Shelley RM and PT Lehtinen (1999) Diagnoses synonymies and occurrences of the pantropical millipeds Leptogoniulus sorornus (Butler) and Trigoniulus corallinus (Gervais) (Spirobolida Pachybolidae Trigoniulinae) Journal of Natural History 33 1379ndash1401 httpdxdoiorg101080002229399299932

Shelley RM CT McAllister CM Nagy ME Weckel RG Christie P Wilson and A Wilson (2011) Distribution of the American milliped genus Boraria Chamberlin 1943 Introductions of B stricta (Broumllemann 1896) in New York and B infesta (Chamberlin 1918) in Connecticut indigenous occurrence of B profuga (Causey 1955) in Louisiana (Diplopoda Polydesmida Xystodesmidae) Insecta Mundi 0194 1ndash8

Shelley RM GB Edwards and AE Bogan (2012) First recorded introduction of the millipede order Stemmiulida (Eugnatha Nematophora) Potential establishment in Florida USA and new records from Mexico northward range extension into southern Tamaulipas Insecta Mundi 0245 1ndash5

Silvestri F (1897) Systema Diplopodum Annali del Museo Civico di Storia Naturale di Genova 38 644ndash651Silvestri F (1903) Classis Diplopoda Vol 1 Anatomie Pars I Segmenta Tegumentum Musculi Acari Myriapoda et

Scorpiones huscque in Italia reperts 346 1ndash272Veacutelez MJ (1967a) New records of Puerto Rican Diplopoda with note of their geographic distribution Stahlia 8 1ndash11Veacutelez MJ (1967b) New species of Diplopoda from Puerto Rico with notes on their geographical distribution and ecology

Caribbean Journal of Science 7(1ndash2) 23ndash39

APPENDIX New platyrhacid records from the Western Hemisphere

Herein we provide full localities and sampling data for relevant unpublished familial records that help document the complete range Determinations even to genera are not germane to this objective and samples have not been examined to this degree Some from along the Javari River Peru constitute near-topotypes of Platyrhacus javarianus Schubart 1950 as DMT will confirm and this binomial may also apply to samples from Amazonas DeptEst in both Colombia and Brazil ldquoRMSrdquo represents samples collected by the first author Missing data was not provided on vial labels

CENTRAL AMERICA

Costa Rica Alajuela Prov Salto La Paz Angel Falls 4 9 July 1964 RE Woodruff (FSCA) Cartago Prov Juan Vintildeas April 1903 OF Cook (NMNH) and Turrialba August 1964 (VMNH) Guanacaste Prov SE slope Volcan Miravalles nr Rio Naranjo 2 28 March 1973 DR Whitehead (NMNH) Heredia Prov Waldboden ldquoFarm Hamburgrdquo 11 February 1934 F Nevermann (VMNH) Finca La Selva 4 km (25 mi) SE Puerto Viejo de Sarapiqui 2 20 July 1976 P Calabi (VMNH) 1ndash15 October 1981 CE Griswold (CAS) and along road from La Selva Biological Station and Rio Sarapriqui 9 March 1981 RE Ashton (NCSM) and Finca la Tigra La Virgen 2 6 February 1976 and 12 February 1980 AM Young (MPM) Limon Prov Cairo 2 11 March 1937 HF Loomis (FSCA) Finca Lalola 3 3 3 December 1963 DG Huckaby (FSCA) Cohuita 30 June 1964 DG Huckaby (FSCA) Colonia ITC de Cariari 2 2 2 March 1967 AL Gardner (FSCA) Squirres 21 September 1976 ES Ross (CAS) La Suerte Biological Field Station August 1979 CL Hogue (LACM) and Suretka 2 (NMNH) Puntarenas Prov Golfito 30 October 1950 ES Ross AE Michelbacher (CAS) 28 km (175 mi) SW Rincon de Osa Rio Agua Buena 2 5 20 June 1964 DG Huckaby (FSCA) 21 km (131 mi) S San Isidro del General 21 June 1964 RE Woodruff (FSCA) Dominical 27 July 1968 M Rivera (CAS) and Monteverde Forest Preserve 40 km (25 mi) N Puntarenas and 25 km (156 mi) NNE Pan American Hwy 4700rsquo 23 April 1963 RO Arnold (FSCA) along Rio Gaucimal 4200rsquo 23 April 1967 (FSCA) 4 5 19 March 1973 DR Whitehead (NMNH) 12 August 1976 P Calabi (VMNH) 17 September 1976 ES Ross (CAS) and 1400ndash1700 m mountain bald 2 10 March 1981 RE Ashton (NCSM)

Panama Chiriqui Prov Reserva Fortuna Continental Divide 4 1 June 1994 NJ Smith R Kassabian (NCSM) PanamaProv Cerro Azul Los Altos 24 May 1994 NJ Smith R Kassabian (NCSM)

SOUTH AMERICA

Brazil Amazonas Est 3ndash5 km (22 mi) E Leticia Colombia in logs in jungle 8 July 1977 RMS (NCSM) and near Marco under log on farm 11 July 1977 RMS (NCSM)

Colombia Amazonas Dept ca 2 km (125 mi) N Leticia along roads to lakes behind airport 10 July 1977 RMS (NCSM) 20 km up Amazon R from Leticia 3 km (19 mi) up Arara Quebrado Ticuna Indian Village 14 July 1977 RMS (NCSM) and 30 km (188 mi) up Amazon R from Leticia 1 km (06 mi) up Tucuchira Quebrada Yagua Indian Village 14 July 1977 RMS (NCSM) Valle Dept 9 km (56 mi) W El Salado in dense forest near small tributary of Rio Anchicaya 2 Smalley Zapata (FSCA)

Ecuador Los Rios Prov 16 km (10 mi) N Quevedo 3 February 1966 R Gilbert (FSCA) Napo Prov N of Tena 12 September 1990 RW Bouchard (NCSM) Pichincha Prov 46 km (288 mi) E Alluroquin 10 April 1984 RW Sites (NCSM) Santa Domingo de los Tsaacutechilas Prov 8 km E Santo Domingo (de Los Colorados) 10 May 1990 NJ Smith (NCSM) and 12 km (75 mi) E Santo Domingo (de Los Colorados) 15 May 1985 C Cervenko (UMSP) SucumbiacuteosProv Limoncocha Biological Preserve 2 15ndash30 October 1964 PW Harman (FSCA)

Peru Loreto Reg Yanomono Kuehn Amazon Expedition 2 14 September 1974 PA Holzbauer (MPM) Moropon Kuehn Amazon Exp 26 October 1974 PA Holzbauer MPM) along Yavari R ca 795 km (497 mi) upstream from Benjamin Constant Amazonas Est Brazil under log in abandoned field RMS (NCSM) along Yavari R ca 80 km (50 mi) upstream from Benjamin Constant Amazonas Est Brazil in abandoned field 20 July 1977 RMS (NCSM) along Yavari R 815 km (509 mi) upstream from Benjamin Constant Amazonas Est Brazil in log at small farm 19 July 1977 RMS (NCSM) along Amazon R 47 km (294 mi) downstream from Iquitos Explorama Lodge 9 August 1986 MA Brittain (NCSM) and along Amazon R 50 km (312 mi) downstream from Iquitos 2 23 July 1993 MA Brittain (NCSM) Pasco Reg Pan de Azuca July 1961 FS Truxal (LACM)

Key to species

Proaspis aitia Loomis 1941

(Hoffman 1980) familial taxonomy (re)elevating Euryurinae and Aphelidesmini to familial ranks with Amplininae remaining a subfamily of the latter Platyrhacidae was left with seven tribes but no subfamilies so we elevate all present tribes (Hoffman 1998 2001a Jeekel 2007) these nine representing new statuses Barydesminae Diodontodesminae Erythrhacinae Hoplurorhachinae Phyodesminae Polydesmorhachinae Psammodesminae Psaphodesminae and Taphodesminae

The Antillean island arc of the eastern and northern Caribbean Sea harbors a curious and spotty mix of what we consider indigenous polydesmidans Of the four suborders (Shelley 2003a Shear 2011) Dalodesmidea is absent and Polydesmidea is widespread being known from all the Greater islands (Cuba Hispaniola Jamaica Puerto Rico) and all the Lessers that have been investigated plus the Bahamas (New Providence) which are actually in the Atlantic Ocean Antigua Dominica Grenada Grenadines (Carriacou) Guadeloupe (Basse-Terre) Marie Galante Tobago St Eustatius St Kitts St Lucia St Martin St Vincent and US Virgin Islands (St John) (Chamberlin 1918a Loomis 1933 1934 1936 1937 1938 1941a 1970 Veacutelez 1967a b de la Torre y Callejas 1974 Mauriegraves 1980 1988 Gonzalez Oliver and Golovatch 1990 Hoffman 1999 Peacuterez-Asso and Peacuterez-Gelabert 2001) Indigenous representatives of Leptodesmidea and Strongylosomatidea however occur sporadically particularly in the Lesser Antilles (Fig 1) Only one potentially native strongylosomatidean is known Mestosoma semirugosum (Pocock 1888) on Dominica (Pocock 1888 Chamberlin 1918a Hoffman 1960a 1999) and the last author suggested that it was introduced from an unknown source area Shelley and Golovatch (2011) however considered it indigenous and reflecting occurrence on the ldquoproto-Antilleanrdquo terrain before it rifted from ldquoproto-South Americardquo in the CretaceousPaleocene with isolation on present-day Dominica when the arc subsequently fragmented Antillean leptodesmideans comprise three families (Fig 1) Chelodesmidae in the Bahamas all the Greaters and Grenada and St Vincent Sphaeriodesmidae in Cuba Hispaniola (Dominican Republic Haiti) Jamaica and Guadeloupe (Basse-Terre) and Platyrhacidae in Hispaniola (Haiti) Guadeloupe (Basse-Terre) and St Lucia (Chamberlin 1918a Mauriegraves 1980 1988 Jeekel 1980 Hoffman 1980 1999 Shelley 2003b 2007 Peacuterez-Asso and Peacuterez-Gelabert 2001 Peacuterez-Gelabert 2008) As specified by Hoffman (1999) Hispaniolan sphaeriodesmids and platyrhacids are known only from Haiti and not from the adjacent and more heavily forested Dominican Republic Here we address Antillean platyrhacids which plausibly also arose on the ldquoproto-Antilleanrdquo terrain and became isolated on their present islands after rifting and fragmentation although Hoffman (1980 1999) and Jeekel (2007) thought that Nannorrhacus may represent a human introduction from the Indomalay Region Similar in their small body-sizes Antillean platyrhacids possess dissimilar gonopods and can only be interpreted as representing distinct monotypic genera

The first mention of a potential West-Indian platyrhacid was by Bollman (1888) who described Stenonia maculata from Cuba and commented that it ldquobelongs in the subgenus Stenonia (= Platyrhacus)rdquo Chamberlin (1918a) included ldquoPlatyrhacus () maculatusrdquo in his list of West Indian myriapods but Loomis (1950) and Hoffman (1999) referred the milliped to Pyrgodesmidae and the genus Docodesmus Cook 1896 Pocock (1894) described Platyrhacus luciae from St Lucia and Cook (1896) designated it type of the new genus Nannorrhacus Chamberlin (1918a) and Hoffman (1999) recognized P and N luciae respectively Hoffman (1960b) claimed the latter had been ldquofairly well describedrdquo but not in accordance with 21st century standards and Jeekel (1980) illustrated a gonopod and paranotum Loomis (1941b) proposed Proaspis aitia n gen n sp for three female platyrhacids in the southern peninsula of Haiti which was subsequently cited by Jeekel (1963) Hoffman (1999) Peacuterez-Asso and Peacuterez-Gelabert (2001) and Peacuterez-Gelabert (2008) Jeekel (1980) described and illustrated Proaspis sahlii from Basse-Terre Guadeloupe that Mauriegraves (1980) also illustrated while adding 25 new localities Though listing P sahlii Hoffman (1999) observed that congeneric status with P aitia could not be assumed until male topotypes of the latter became available and that acropodal differences between Jeekelrsquos and Mauriegravesrsquodrawings done from closely similar perspectives ldquostrongly suggest that two species are involvedrdquo

While perusing millipeds at the FSCA (acronyms below) in 2011 RMS discovered two platyrhacid samples one with five males collected in 1984 at Morne Formond a small mountain in Parc National de Macaya Sud Department Les Cayes Arrondissement near the western end of the southern Haitian peninsula The site is around 8ndash16 km (5ndash10 mi) SW of the type locality an unspecified site between Camp Perrin a municipality in the same arrondissement and department and Riviegravere Glace in GrandrsquoAnse Department We consider this sufficiently proximate to constitute near topotypes of Proaspis aitia particularly with an indefinite type locality and without other males Seventy-one years after its proposal we therefore establish the identity of P aitia and describe and illustrate males Since its biramous gonopodal telopodite is distinct from the uniramous one of P sahlii we propose

Key to species

Conclusion

Acknowledgments

Literature cited

CENTRAL AMERICA

SOUTH AMERICA

Key to species

Conclusion

Acknowledgments

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CENTRAL AMERICA

SOUTH AMERICA

Key to species

Conclusion

Acknowledgments

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CENTRAL AMERICA

SOUTH AMERICA

Key to species

Conclusion

Acknowledgments

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CENTRAL AMERICA

SOUTH AMERICA

Key to species

Conclusion

Acknowledgments

Literature cited

CENTRAL AMERICA

SOUTH AMERICA

Key to species

Conclusion

Acknowledgments

Literature cited

CENTRAL AMERICA

SOUTH AMERICA

Key to species

Conclusion

Acknowledgments

Literature cited

CENTRAL AMERICA

SOUTH AMERICA

Key to species

Conclusion

Acknowledgments

Literature cited

CENTRAL AMERICA

SOUTH AMERICA

Key to species

Conclusion

Acknowledgments

Literature cited

CENTRAL AMERICA

SOUTH AMERICA

Key to species

Conclusion

Acknowledgments

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CENTRAL AMERICA

SOUTH AMERICA

Key to species

Conclusion

Acknowledgments

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CENTRAL AMERICA

SOUTH AMERICA

Key to species

Conclusion

Acknowledgments

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CENTRAL AMERICA

SOUTH AMERICA

Key to species

Conclusion

Acknowledgments

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CENTRAL AMERICA

SOUTH AMERICA

Key to species

Conclusion

Acknowledgments

Literature cited

CENTRAL AMERICA

SOUTH AMERICA

Key to species

Conclusion

Acknowledgments

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CENTRAL AMERICA

SOUTH AMERICA

Key to species

Conclusion

Acknowledgments

Literature cited

CENTRAL AMERICA

SOUTH AMERICA

Key to species

Acknowledgments

Literature cited

CENTRAL AMERICA

SOUTH AMERICA

Key to species

CENTRAL AMERICA

SOUTH AMERICA

Key to species

CENTRAL AMERICA

SOUTH AMERICA

Key to species

CENTRAL AMERICA

SOUTH AMERICA

Key to species

SOUTH AMERICA

Key to species

Documents

Notes on a collection of Myriapoda from Cuba