Great Basin Naturalist 53(3),259-281

REVISION OF THE GENERA OF PLATYPODIDAE (COLEOPTERA)

Stephen L. Wood l

AIJ~lllAcr.- A search for characters to make the generic classification of Platypodidat: more nearly ohjective result­ ed in the discovery ornew anatomical fcahnes tflat appear to reneet p<1Uerns in phylogeny for this family. The PlaLypo­ didae are members of the Curculionoidca. and are very closely allied to Scolyticlae. Three subfamilies of Pl.ltypodidae are reco~'11ized: (1) C'.optonotinae, containing; Coptonotini (Coptonotus Chapuis, Protohylastes Wood, Sw[.ytowrsus School), Mecopelmini (Mecopelnws Blackman, Protoplatypus Wood), nnd Schedlarini (Schedlarius Wood [=CJuqmisia Du?;es)); (2) Tesserocerinae, containing Diapodini (DUlpUS Chapuis, Cenyocerw Motschulsky [=Dincavus SchcdlD. and Tesseroccrini (Platytarsu/ns Schedl, Notoplnt.ypU$ Lea. TesserocranuLus Schedl, l'esserocert15 Saunders [=Damicerus Spinola, TesseroplalljpU-'t Schedl, Tesserocephalw Schedl}), Spathidi.ce11l$ Chapuis. l'eriommatus Chapllis [=Asettls Nunbcrg, Setatlus Nllnbergl, Chaetustus Nllnberg [=Sym11U--'l"us Chapuisl, C'-"TUxephalu$ Chapnis, and Mito$oma Cha­ puis [=Platypicerus NllIlberg, COtJcephalonus Sched!}); and (3) Platyplxlinae, containing Platypodini (l'latYrnls Herbst [=Cylindra T1liger, Steno1'Jlalypus StrohIlieyer, Platypinus Schecl!, Atl.Stroplaiypus Brow!l{~J, Treptopl(lt!lpus Schecl!, Grossotar8fl.S Chapuis. Trachyost/,ls Schedl, Neotrachyostus Browne, Platyscaplllw Schcdl [=PlatyscupU.8 Schcdl, Costaruplotus Nlmbel'ro, Baiocis Browne, Cylindropalpus Strohmeyer, TrWuwtus Scht::dl, MeSU1J{m,ypuS Slrohmeyer, IJoliorygt.l$ Schcdl (=SCUWTJygll·' Nunberg, Pygudolim Nunberg, Mixopy~us Nunberg, Mesopygus Nlmhergl. and 11 genera named as new to scien~ derived from the g;enus Platypus of S(:hedl 1972. The following are new genera and their typc·specics: Peroplatypu,Y (for Platypus truncatipennis Sehedl), Dinoplt.ttypus (fl)r Plmypus Cupula/Uk Chapuis), MyvplatyptJS (for Bostrichus or P/atypw jlaui.comis Fabl'iLiHS), Oxvplat!fPUS (for Scolytus or Platypus C{UO(lritk."ntatus Olivier), Platyplu;sus (for Platypus obtusus Chapuis), MegapultYPU8 (for Platypus dentaLus Dalman), Euplntypus (for BObtrichw or Platypus parallelu.s Fabricius), Epiplatypus (for PlalYTJUS (~'ceptor Wood). and TeloplatypU$ (fol" Platypus concinnus Blandford). lne archaic "seklionen" u~ by Schedl ill his 1972 classification ofthis ramily are eliminated. A key for the idenlifiC'atlon orgenera, a dhcussioll ofcharacters, ami remarks on phylogtlny arc included.

Key words: P!4lypodidae, Coleoptera, Platypus, n:vision, taxonomy.

During prepartion ofthe recent world cata­ now contains slightly over 1400 species, log that included the family Platypodidae almost aU ofwhich arc tropical in distribution. (Wood & Bright cl992), it was leamed with By 1864 approximately 17 species had considemblc surprise that a systematic review been assigned to Platypodidae in Pkltypus, of genera for this family did not exist. This Tesserocerus, and ee"!loeerus. Chapuis (1865) contribution is written in an attempt to at added scven genera (Cetwcephalus, Grossotar­ least partially remedy that situation. sus, Diapus, Mitosomu, Periommatus, The group had its origin in systematics lit­ Spathidicerus, Symrnerus [=Ghaetaslus)) and erature when Fahricius (1792:364) named about 220 species to the family in his classical Bostrichus cylindrus from Germany and Monographic des Platypides. This monograph assigned it to the non-Linnaean Xylophaga in (Chapuis 1865:22-23) contained the first key the family Bostrichidae. A year later Herbst to genera used in the family. It was based (1793: 128) recognized the uniqueness of this largely upon mouthparts, eyes, and features of spccies and erected the genus Platypus for it. the prothorax. Species in the larger genera Platypus was transferred by Latreille were divided into several archaic. non-Lin­ (1807:277) to his newly erected subfamily naean species-groups that were perpc.hlated Scolytarii (cunently Scolytidae) of his family by Strohmeycr (1912, 1914b), Sehedl (1939, Curculionites. Sbuckard (1840 [reprinted 1972) and, to a lesser extent, by other authors. 1861:64]) estahlished the family Platypodidae Strohmeyer (1912) broadened the family to for it. The group has received a tribe, subfam­ include the subfamily Chapuisiinae for Cha­ ily, or family designation in virtually all treat­ puisia Dug,,, (=Schedlarius), hut he later ments of the group since 1840. The family placed it in a separate family, Coptonotidac

.l3J2 urc SelellCC Muscum.nrigh~m Young Univcrsily.1'Tovo. Utah ISo4602.

259 260 GHEAT BASI~ NATURALIST [Volume 53

(Strohmeyer 1914a), a change followed only While attempting to organize a reasonable by Schedl (1939). Strohmeyer (1914b:18) arrangement of genera for the world catalog, I divided the 323 known species of Platypodi­ obselVed that some obviously related species dae into two subfamilies based on the divided had been grouped by Schedl (1972) in entirely (Tesserocerinae) or undivided (Platypodinae) different subfamilies, while other unrelated maxilla. The Tesserocerinae he then divided species had been clustered into the same into trihes Tesserocerariae (Tesserocerus, Peri­ genus, and I recognized that a serious generic ommatus, Spathidicerus), Symmerariae (Sym­ revision has not been puhlished on this taxon mems), Cenocephalariae (Cerwcephalus, Mito­ since the family was first recognized. The soma), and Oiapodariae (Diapus) based on archaic classification then in use was unrea­ procoxal, oecuIar, and funicular characters. sonable, unwieldy, and based as much on the The Platypodinae were divided into tribes whims of the taxonomist using it as on phy~ Platypodariae (Platypus, Cylindropalpus, logeny or evolutionary relationships of the Notoplatypus) and Crossotarsariae (Crossotar­ included taxa. Tbis led to a search for charac­ sus, Stenoplatypus, Mesoplatypus) based on ters that might be usable in a new classifica­ characters ofabdominal sternum 8. Strohmey­ tion. er perpetuated and expanded the non-Lin­ naean species-groups of Chapuis in his cla'lsi· REVIEW OF CHARACTERS fication of the larger genera. Schedl (1939) proposed a superfamily The Platypodidae are members of the Scolytoidea in which he placed the families superfamily Curculionoidea (Crowson 1955, Scolytidae, Coptonolidae (for Coptonotus, 1968). They share many characters with other Scolytotarsus, Chapuisia), Platytarsulidae (for members of this group (Wood 1973, 1986). Platytarsulus, Notoplatypus), and Platypodi­ Within the Curculionoidea tbey are very dae, with no subfamilies indicated, containing closely allied to the Scolytidae with which tribes Platypodini, Tesserocerini, Ceno­ they share the same broad ecological niche cephalini, Crossotarsini, Periomatini [sic], and and many similar behavioral patterns. Togeth­ Oiaporini [sic]. Schedl's (1972) Monographie er these two families share a conspicuous del' Familie Platypodidae elevated the Crosso­ pregular sclerite (as defined by Hopkins 1909, tarsinae, Platypodinae, Periommatinae, and 1911) that is clearly marked by sutures on Diaporinae [sic] from tribal to subfamily rank, both sides (Figs. 3-5) and is not similarly marked in any other family ofCurculionoidea. but reduced to subfamily rank the Platytar­ Additional characters shared by these two sulinae. The Tesserocerini and Cenocephalini families and those features that distinguish were grouped within his Platypodinae. Schedl them from allied families are reviewed by (1962) treated in his family Coptonotidae the Wood (1973, 1986). The featore most familiar genera CoptallotUS, Schedlarius (=Chapuisia), to coleopterists and the one most widely and Mecopelmus. In his treatment of Platypo­ employed in family keys for separating Platy­ didae, Schedl (1939, 1972) perpetuated the podidae from Scolylidae is the length of tarsal use of the non-Linnaean species-groups of segment 1 compared to segments 2-5. In Chapuis with only minor modifications. Sc'Olylidae segmellts I, 2, and 3 are su bequal Wood (1973, 1986) included the Coptonoti­ in length, while in Platypodidae segment 1 is nae (Coptonotini, Mecopelmini, Schedlarini) usually about as long as segments 2-5 com­ in P]atypodidae. Wood (c1992 ill Wood & bined (Wood 1986:11, fig. 9). However, in Pro­ Bright) recognized the subfamilies Coptonoti­ toplatypus (Mecopelmini) segments I, 2, and nae (tribes Coptonotini, Mecopelmini, Sched­ 3 are subequal in length, while in Mewpelmus larini). Tesserocerinae (tribes Diapodini, (Mecopelmini) segment 1 is as long as 2-5 Tesserocerini), and Platypodinae (tribe Platy­ combined. The remaining species of podini). A dendrogram tbat indicates possible Coptonotinae are intermediate in their phylogenetic relationships among these ex'Pression of this character (Schedl 1939:381, groups to one another and to Scolytidae fig. 3). appears in Wood (1982:43), except that the The pregula ,clerite in Coptonotinae (Fig. Tesserocerinae and Platypodinae are not 3) is comparatively small as it is in Scolytidae; divided. in Tesserocerinae (Fig. 4) and Platypodinae 1993J REVISION OF PLATYPODIOAE 261

1 "/ 2 '" .... '7 , I '\ /' , ~ , ~ 0/

4

3 5 ..-.. -. -

Figs. 1-5. Platypodidac spp., males: 1, Schedlarius mexicaTlus (Chapuis), lateral aspect ofthorax: showing ::;traight pos­ terior margin of prothomx (upper arrow) and anapleural suture of mesothorax (lower arrow); 2, l'esserocerus dewalquei Chapuis, lateral aspect of thorax showing strongl)' pl'Ocurved posterior margin of prothorax (arrow) and absence of a mesothoracic anaplcural suture; 3, Schedlarius mexicanus, ventral aspect ofhead showing the small pregula (arrow) with its marginal sutures and transverse carina; 4, Tessuocerus d<'W~alquei, ventral aspect of head showing the deep cleft (stippled area at arrow) between the large pregula and margin of the ordl fossa; and 5. Euplatypu$ para11elw (Fabricius), ventral aspect ofthe head showin~ absence ofthe deft (arrow).

(Fig. 5) it is much larger and very conspicu­ and galea lobes (Wood 1986:8, fig. 6); its ous. In Tesserocerinae there is a conspicuous absence appears to be correlated with the cleft (Fig. 4, arrow) between the lateral margin fusion of tbe lacinia and galea into one ele­ of the pregula and the margin of the oral fossa ment. Due to Ihe paucity of specimens avail­ (into which the maxilla moves). The depth of able for study, Platytarsulus and Notoplatypus this cleft is usually equal to at least half the were assigned to Tesserocerinae on the basis length of the pregula (Fig. 4). In Platypodinae of the presence of the cleft and were not dis­ this cleft is very shallow to nonexistent and is sected to detennine tI,e character ofthe maxilla. always equal in depth to less than one-fourth In the Tesserocerinae the eye exhibits a the length of the pregnla (Fig. 5). The pres­ departure from the usual subcircular, hemi­ ence of this cleft is apparently correlated with spherical shape. [n Platytarsulus, Spathidicerus, the division of the maxilla into separate lacinia and Periommatus the eye may be very large 262 GREAT BASIN NATURALIST [Volume 53 and reniform (Schcdl 1939:384, fig. 4). In ed in numher. A feature, apparently used here Mecopelmm (Coptonotinae) there is a slight for the first time, is the presence of a groove modification in that direction (Blackman or impression on the posterior portions of the 1944:figs. 3-5). metasternum and metepisternum for the Antennal characters appear to be signifi­ reception of the metafemur. In the African cant in the emly phyletic history of the Platy­ genera Cylindropalpus, Triozastus, Mesoplaty­ podidae. The antcnnal club is weakly marked pus~ and Doliopygus, the anterior margin of by two strongly procurved sutures in this impression is continuously carinate (or Coptonotus; in the remainder of thc family nearly so). In more than half (mostly Amcrican there is no evidence of sutures on the club. species) ofwhat has previously been designat­ Coptonofus has the antennal funicle 7-seg­ ed as Platypus, the anterior margin of this mented (Schedl 1939:380, Hg. 2), a feature impression is marked by a series of minute also shared with Protohylastes and Scaly totar­ spines, and in the remaining half of Plntypus $US. Because seven is the maximum number of (mostly African and Indo-Australian species) segments in the funicle found in Cur­ the impression is weak to absent and spines culionoidea (Crowson 1955, 1968), that num­ are absent. Except for females ofa few Ameri­ ber is assumed to exhibit the primitive charac­ can species, this character appears to be a ter; any reduction from that number should reliable indicator ofrelationship. represent specialization. In Protoplntypus and 11,e visible abdominal sterna in Platypodi­ Schedlarius the funicle is 5-segmcnted, in dae exhibit rather limited, but remarkable, Mecopelmus and Notoplntypus 3-segmented, variation. The primitive structure appears to in Pl4trjtarsulus 2-segmented, and in all other be five horiwntal. unarmed segments that rise Tesserocerinae and in all Platypodinae it is 4­ little, if any, to meet the apex of the elytral segmented (Schedl 1939:380, fig. 2). In declivity. As the male declivity shortens (or Coptonotus and Protohylastes the antennal atrophies) in some groups, the ahdomen club is more slender and less strongly flat­ ascends gradually to abruptly to accommodate tened than in other representatives of the the change. [n Diapodini (Diapus, Genyoeerus) family. this ascent is almost entirely accomplished by In Coptonotinae the posterior margin of visible sternum 5 as it becomes vertical and tl,e prothorax (as seen from a lateral aspect) is moderately to remarkably concave (This dorsoventrally straight to very weakly enables males of these genera forcefully to procurved; the mesepisternum is moderately expel frass from the entrance hole 2 m or large and almost flat (Fig. 1). In Tesseroceri­ more from the host tree). In Mesoplatypus and nae (Fig. 2) and Platypodinae the posterior Doliopygus a pronounced transverse carina margin of the prothorJ.x is strongly procurved occurs on visihle sternum 2 (SchedlI972:149, in the pleural area, the mcsepisternum is fig. 39); this enables stcrna 3-5 (at least in inflated, or in specialized genera it may be Doliopygus) to become vertical and concave impressed and variously ca.-inate. and to function much as does sternum 5 in the In all Coptonotinae and in Diapodini, Diapodini. Less remarkable and less extensive Tesserocerus, and Tesserocranulus of the variations occur on sterna 3, 4, or 5 where a Tesserocerinae, the mesotergum is normal, transverse pair of moderate to elaborate that is, transversely flat or weakly, transversely spioes (Wood 1966:47 [fig. 6], 67 [Hgs. 22-24]) convex. In all other Tesserocerinae and all arm one of these segments (sternum 3 in Platypodinae it is armed by a conspicuous, Myoplntypus, sternum 4 in Oxypl4typus, ster­ strongly elevated, median carina. The pres­ num 5 in Plntysearulus). ence of this carina represents an obvious spe­ The protihia is somewhat uniform in the cialization. In ScoJytidae and Coptonotinae family except in primitive genera. It charac­ the mesothoracic anapleural (pleurosternal) teristically ha.o;; a terminal mucro and is :u·med suture is consistently present. This anapleural on the posterior (or lateral) face by one or suture is largely (lbsserocerinae) or entirely more transversely carinate, coarse rugae. In (Platypodinae) eliminated in the higher Platy­ female Crossotarsus, Trachyostus, and Neotra­ podidae. chyostus, and in at least two species of Ameri­ Characters that might be used to indicate can Megaplatypus, these rugae are broken up phylogenetic trends in Platypodinae are limit- and reorganized into numerous, confused 1993] REVISION Or. PLATYPODlDAF. 263 granules. The socketed denticles (derived These patterns appc..ar morc constant and less from setae) found in most Scolytidae (Wood diverse than previous usage might suggest. 1986:11, fig. 10) are unknown in Platypodidae. The three genera of Coptonotinae studied The tibial denticles of Platypodidac are true in the field by me have habits more nearly like spines that nmetion in gripping tunnel walls. Scolytidae than like other Platypodidae. In The simplest fi,rm appears to be that of Proto­ Schedlarius (Wood 1957), parent adults make hylastes (Wood 1973:86, fig. 25). Other long egg tunnels in the xylem; egg niches are Coptonotinae may have one lateral spine or randomly formed on al1 sides of the tu nncl carinate ruga; higher genera have two to nine into which the eggs are placed, onc in each rugae (Wood 1973:86, figs. 25-33, Schedl niche, and packed in frass. The larvae form 1939:379, fig. 1). The rugae are not always long, independent, winding tunnels in the consistent in position and form in the higher xylem. Altbough fungal decay in tbe vicinity genera and must he used in classification with of boring activity of Loth adult" and larvae was caution. ohvious, there was no ambrosial mycelium Tarsal segment 3 is slender and cylindrical growth on the waJls of adult or larval mines. in almost all Platypodidac (Wood 1986:11, fig. Adult Mecopelmus form a simple cave tunnel, 9). In the Coptonotinac genera Coplonolus, about 1 em in diameter and 1-2 mm deep, at Prulohylastes, Scolylou,rsus, and Sclwdlar'ius and slightly below the cambium region of segment 3 (Schedl 1939:381, fig. 3) is broad their host. In the frass of this chamber the and strongly bilobed as in primitive Cur­ female deposits a cluster of one or two dozen culionoidea. ew. rl11e first- and second-instar larvae feed The spines arming the male clytral declivi­ on this Ji'ass and then bore individual mines in ty are truly remarkable and almost endlessly the cambium region as they radiate out from diverse within the Platypodidae. However, as the central chamher. There was no evidence groups are segregated on the hasis of other of mycelial growth in the mines at 20X magni­ characters, the constancy and usefulness of fication. Protoplalyp1ls parent adults formed these spines and patterns of spines become apparent. Greater knowledge of Platypodidac radiate tunnels (with 3-5 egg galleries in each, simjlar to those of in the cam­ pairing and mating behavior would probably Pilyophthorus) increase our understanding of the significance bium of thei.. host, complete with nuptial ofsize and posil:ion of these spines. chamber, egg galleries, egg niches, and inw­ It is generally underslood that all Platypo­ vidual larval mines. Mycelial g:rowth was not didae (except Protoplatypus) are monugynous, evident at 20X magnification. All other and in all species the male initiates the forma­ observed Platypodidae (Tesscfocerinae and tion of a new parental gallery system. The Platypodinae) are xylomycctophagolls. female, consequently, aSSumes the primary responsibility for the identification and accep· PIIYLOGENY tance of a male. Presumably, for this reason, the male postedor extremities exhibit remark­ The Platypodidae and Scolylidae are very able characters, while the females exhihit few, closely related to one anothel~ so much so that often very subtle, distinguishing features. Per­ four of the six genera of Coptonotinae could haps the most remarkable female features are be assigned to either family without serious the dehiscent mandibular appendages of the conflict. The most closely allied groups within Diapodini (Roberts 1993) that are used briefly the Seolytidae to the Coptonotinae, however, for tactile communication with the male at appear to be in or near the Carphodicticini or pairing and/or mating ami a.re then discarded. possibly the Dryocoetini of the Scolytidae, A few other females that possess remarkable rather than the superficially similar tribes in frontal characters apparently use those fea­ what has been re~arded as the more primitive tures in caring fi)r the eggs or larvae (Wood Hylcsininae. The Platypodidae appear to be 1986:4, fig. 2). This is an area where very lillie the first of eight phyletic lines within the factual information is availahle. The mycetan­ Platypowdae-Scolytidae group to adopt the gia pores on the pronotum of many female xylomycetophagous habit. This shili in adap­ (and a few male) Platypodidae appear to fol­ tive specialization had a profound impact on low distinguishable patterns in some groups. form and function within the Platypodidae 264 GREAT BASIN NATURALIST [Volume 53 that separated them rather sharply from the female Tesserocerini has not been clarified. Scolytidae. Evolution within the Curculionoidea is obvi­ Few detailed anatomical studies have been ously much more complex than publislled based on members of the PJatypodidae, and, simplistic explanations acknowledge. Obvi­ for the most part, those that have been made uusly many unanswered questions remain that were based on the accessible, more special· must be 'IDswered before reasonable explana­ ized representatives. tions are found. It is almost universally agreed that the To summarize the above, it appears that the Platypodidae-Scolytidae are members of the Platypodidae-Scolytidae represent a distinct Curculionoidea (Crowson 1955, 1968:154-166, phyletic line of Curculionoidea having; one Wood 1973, 1986). Exactly where these fami­ gular suture and that this line is independent lies fit within the Curculionoidea has been the from the Brenthidae-Rhynchitidae-Curculion­ subject of much discussion and disagreement. idae line ofspecialization. Their traditional placement within (Crowson Phyletic trends within the Platypodidae are 1955, 1968, as subfamilies) or adjacent (as somewhat unclear. The six genera of families) to Curculionidae is questioned by Coptonotinae (represented by nine rare me (Wood 1973, 1986). Their placement species scattered on four tropical contenents among the higher Curculionoidea is substanti­ and New Guinea) appear to represent relict ated by the presence of only one median, remains of a ooce much larger group. All lack gular suture (Wood 1986:6, 8); bowever, the the median mesonotal carina once thought to very short length of this suture in Platypodi­ characterize all representatives of this family, dae is a departure from most other Cur­ and all have the anapleural suture on the culionoidea and could have significance. The mesopleuron. Four of these genera (CopI01l0­ comparative positions of the mandibular tus, Protohyl..,·ws, Scolytotarsus, Schedlarius) condyles, including conspicuous reduction of have tarsal segment 3 broad and bilobed. the hypostomal area, also sets the Platypodi­ None of the six has the antennal funicle 4-seg­ dae-Scolylidae apart from other Cur­ mented as it is in all but two genera of the culionoidea, particularly the Cossoninae (Cur~ remainder of the family. However, none of culionidae) to which they are supposed to be these six genera has the maxilla divided into closely related (Wood 1986: lo, fig. 8). The sepamte Jacinia and galea. The three genera truly unique character of the Platypodidae­ for which habits are known all lack the xylo­ S<.'olytidae is the conspicuous pregular sclerite cycetophagous habit. Of these six genera, that is clearly marked on both sides hy sutures Coptonotus and Protohylastes are closely (Figs. 3-5), a feature that is shared by no other allied to one another and approach the family (Wood 1986:6-8). In Anthribidae and Scolytidae more closely in structural detail Nemonychidae the lacinia and galea form sep­ than do the other fOlll: SchedJ.arWs appears to arate elements on the maxilla. Among those be the most closely allied to other Platypodi­ Curculionoidea having only one gular suture, dae of tlmse six. ProtDplatrjptlS and Mecopel­ the maxilla is similarly divided only in some mus are allied to one another but appear to Attelabidae, some Rhynchitidae, and the sub­ represent an independent evolutionary exper­ family Tesserocerinae of the Platypodidae iment with no close alJiance to any other (Wood 1986:8, fig. 6). No member of the Cur­ group. culionidae shares this character. The loss of The Tesserocerinae are characterized by the mesothoracic anapleural suture in Tesse­ the division of the rnaxiUa into separate lacinia I'ocerinae (Fig. 2) and Platypodinae appears to and galea elements (Wood 1986:8, fig. 6) and have occurred entirely within the Platypodi. by the accompanying cleft between the pregu­ dae because it is present in aU Coptonotinae la and margin of the oml fossa (Fig. 4). This (Fig. 1) and in all Scolytidae. Browne (1972) feature suggests a relationship to the most reported urogomphi-Jike structures in larvae primitive Curculionoidea families. Within the of two species of African Platypodidae; if cor­ subfamily, the Diapodini (Diapus, Genyo­ rect, this would be the only known occurren(,'C cenlS), Tesserocerus, and Tesserocra.rwlus lack of these structures in Curculionoidea. The a median carina on the mesotergum. The true homology of a labrum-like structure in Diapodini also have the procoxae widely sepa­ Chaetastus (Wood 1986:4, fig. 2) and other rated, a primitive feature, but the mycetangia 1993) REVISION OF PLATYPODlDAE 265 pores on the pronotum and the highly special­ recent invasion of Africa, and one species of ized abdominal sternum 5 represent extreme uncertain affinity in tropical America. Group specialization. Platytarsulus (2-segmented) b appears to have occupied the Indo-Aus­ and Notoplatypus (3-segmented) have a tralian, African, and South American areas reduced number of segments in the antennal before the separation of Africa and South funicle. These six genera have the protibia America and suggests a pre-Tertiary origin. more slender and with fewer transverse, cari­ Group c is primarily American except for nate rugae on the posterior (or lateral) face. Baiocis that is of uncertain affinity. and They probably represent the more primitive Euplatypus that appears to have had early element of the family after family characters interaction with Madagascar and a later were firmly fixed. exchange from Madagascar to Africa [The The (a) Diapodini, Platytarsulus, and Noto­ modern circumtropical extension through platypus are exclusively Indo-Australian in commerce of pat-allelus from America is distribution (except for one species of Diapus ignored]. Group d is exclusively African recently introduced through modern com­ except for one species that reached Madagas­ merce into Africa) and each is without a close car from Africa rather recently. Based on these living relative; (b) Spathidiceros (Indo-Aus­ data, it appears that evolution of the Platypod­ tralian) and Periommatus (African) are obvi­ inae has been rapid since the early Tertiary ously derived from a common ancestor and separation of Africa and South America and are closely related to one another; (c) Tessero­ that pre-Tertiary taxa must have been struc­ eerus and Tesseroeraoulus (both tropical turally very different from modern species. American) are also allied to one another; (d) Pre-Tertiary PJatypodinae must have resem­ Chaetastus (African), Mitosoma (Madagas­ bled the Coptonotinae much more than has caran), and Cenocephalus (tropical American) previously been supposed and suggests an ori­ are also allied to one another, but are quite gin no earlier than that of flowering plants distinct from other Tesserocerini. It appears (Lower Cretaceous). that groups a, b, and c have evolved entirely since the early Tertiary separation of Africa SYSTEMATIC SECTION and South America. Only group d exhibits a phyletic imprint of pre-Tertiary development. Because this represents the first real exam­ It is concluded, therefore, that the evolution ination of gene.ic classification in Platypodi­ of the Tesserocerinae has been rapid and that dae since the family was established. some pre-Tertiary representatives of this subfamily radical departures from previous treatments must have been radically different from mod­ are recommended. Foremost among these is em taxa. the abandonment of the archaic practice of In the Platypodinae (Platypodini) four lines employing undefinable species-groups or of development are seen: (a) Platypus Qargely infrageneric groups below the genus level and African to Indo-Australian), Treptoplatypus above the species rank. Schedl (1972) (Indo-Australian, Oriental, NW North Ameri­ employed 62 of these groups in his treatment can), Peroplatypus (Indo-Australian), Dino­ of the genus Platypus. This change made it platypus (Indo-Australian); (b) CrossotarStlS necessary to retrieve a number of generic (Indo-Australian), 1l"achyostus (African), Neo­ names that had previously been placed in syn­ trachyostus (tropical American); (c) Platyscap­ onymy and to name several others. Although ulus (tropical American), Myoplatypus (Ameri­ this will cause some initial confusion, it can), Oxoplatypus (American), Platyphyws should ultimately enhance communication on (tropical American), Megaplatypus (tropical this family. American), EuplatyptlS (mostly American. The treatment of genera following the key some African, Madagascaran), Baiocis (Indo­ is brief except in the tribe Platypodini (sub­ Malayan), Epiplatypus (tropical American), family Platypodinae) because of the significant Teloplatypus (tropical American); (d) Cylin­ changes introduced there. The treatment of dropalpus (mostly African, 1 Madagascaran), the six genera of Coptonotinae is virtually Triozastus (African), Mesoplatypus (African). unchanged from previous usage. The signifi­ DolWpygus (African). Group a occurs primari­ cant changes in Tesserocerinae include (1) the ly in the Indo-Australian area, with slight, transfer ofPlatyta,·sulus and Notoplatwus into 266 GREAT BASIN NATURALIST [Volume 53 this subfamily from Platypodinae, and (2) the Antennal funicle 3--5-segmented, club larg­ restoration of Chaetastus, Cenocephalus, and er, broader, more strongly flattened; profe­ mur stouter, less than 2.0 hmes as long as Mitosoma to full generic rank. wide (except slender in Schedlarius); protib­ Tbis study was based on my personal col­ ia stouter, conspicuously shorter than lection of over 400 species of Platypodidae femur; pregula without a median carina or and my examination of more than 400 other spine; either pale species smaller than 2.0 species. Because approximately half of the mm or very slender 5 known species in the family were not seen by 3(2). Eye very large, flat, subreniform; antennal me, it is obvious that adjustments in the pro­ club inconspicuously marked by two strong­ posals made here will be needed in tbe future. ly procurved, suhangulate sutures, these The monobasic genera Crossotarsinulus weakly indicated by grooves and setae; Schedl (1972:84-87) and Spathicranuloides color almost black; tropical America; 3.2 or Schedl (1972:71) are unknown to me and, con­ 9.0 mm Coptonotus Chapuis sequently, were not included in this study. Eye short, subcircular to oval in outline, less than 1.5 times as long as wide; antennal Key to the Genera of Platypodidae club unmarked by sutures; Africa or Aus- tralia '" "' 4 I. Posterior margin of prothorax (as seen fi'om lateral aspect) straight to weakly procmved 4(3). Eye oval, about 1.5 times as long as wide; in pleural area (Fig. 1); mesepistemum protibia very slender. its apex armed by a moderately large, almost flat; mesotergum small median spine, a minor spine on each flat to broadly, transversely arched, without side near apex; body and pronotum hylesi- a conspicuous, acute, median carina, scutel- nine in form; Australia; 9.8 mm .. hun rising abruptly to elytral ~;urface; prono- ...... , Protohylastes Wood tum never with mycetangia grooves or pores; preguJa small, bearing a transverse Eye, subcircular, hemispherical; protibia cmina; eyes sometimes large, elongate, flat; distorted by a large spine near its middle; antennal scape slender, club-shaped; pro- body and pronotum platypodine; Africa and coxae smaller, usually on middle third of Australia; 4.5-4.9 mm Scolytotarsus Sehedl prosternum length; tarsal segment 1 short (except elongate in Mecopelmus, Schedlar- 5(2). Protibia small, with a terminal mucro and ius); anapleural suture on mesothorax pres- with or without one spine on lateral margin; ent (Fig. 1); xylophagous or phloeophagous; tarsal segment 3 narrow, cylindrical; elytra (Coptonotinae) 2 simple, declivity convex, unarmed; pale species, body less than 2.0 mm; (Mecopel- Posterior margin of prothorax strongly mini) 6 proclltved in pleuml area (Fig. 2); mesepi­ sternum large, usually inflated (concave in Protibia larger, subapically armed on lateral some Tesserocerinae); pronotum often with face immediately above tarsal insertion by conspicuous grooves or pores extending one transverse, coarse, costate ruga; tarsal into mycetangia; mesotergum usually bear­ segment 3 very broad, deeply bilobed; ing a conspicuous median carina (absent in antennal funicle 5-segmented; elytral four genera), scutellum, if present, rising declivity obliquely subtruncate, its margin gradually, usually carinate and apically costate on lower half, armed above by a row pointed; procoxae enlarged, occupying pos­ ofseveral small spines, its face on interstriae terior half of segment; pregula moderately ornamented by numerous small, white to very large, usually flat, never with a scales; Mexico to Panama; xylophagous in transverse carina; antennal scape variously Bursel"a spp.; 4.0-7.0 mm; (Schedlarini) .. modified; eyes usually rounded, hemispher­ ...... Schedlarius Wood ical; tarsal segment 1 always elongate, usu- ally longer than segments 2-5 combined; 6(5). Antennal funicle 5-segmented, posterior anapleural suture on mesothorax largely or face of club glabrous; eye smaller, finely entirely absent (Fig. 2); xylomycetophagous ...... 7 faceted; abdomen horizontal, costal margin of elytra horizontal or descending to apex; 2(1). Antennal funicle 7-segmented, club slender, tarsal segment 1 short, subequal in length to small; profemur more slender, at least 2.6 2 or 3; adults polygynous, parental tunnels times as long as wide, protibia more slender radiate, in cambium, first-instal' larvae form and almost as long as femur; pregula with a individual tunnels in cambium; New Guinea; higher median carina arising from low 1.2--1.5 mm Protoplatypu8 Wood transverse carina and terminating cephalad in a small, blunt spine; species larger than Antennal funicle 3-segmented, club pubes­ 3.5 mm, stouter, darker in color; (Coptano- cent on both faces; eye larger, coarsely hni) , 3 faceted; abdomen distinctly ascending 1993] REVISION OF PLATYPODIDAE 267

behind, costal margin ofe1ytra ascending on weakly armed; pronotum and elytra reticu- apical one-fHih; tarsal segment 1 as long as late , , , 11 2-5 combined; adults monogynous, parental chamber a simple cave, third-instal' larvae Antennal funicle 4-segmented, club either form independent tunnels radiating from pubescent to base or glabrous area much central chamber in Setjania spp.; Panama; smaller (basal onc-fonrth); protibia usually 1.4-1.6 mm Mecopelmus Blackman more elaborately armed 12

7(1). Maxilla with mesal clement clearly divided 11(10). Antennal funicle 2-segmented; eye elon­ into separate laciuia and galea; pregula sep­ gate, reniform, at least 2.0 times as long as arated on each side from margin of oral wide; protihia with one transverse ruga fossa by a deep cleft (into which maxilla above lateral spine; Borneo to Malaya; 4.5 moves) equal to at least one-half pregula lflm jf'ltztytll1",~ttltl.~ 1)clItlcll length, visible pregula caudad from cleft comparatively small; (Tcsserocerinac) 8 Antennal funicle 3-segmented; eye suhcir­ cular, as wide as long, entire; protibia with a Maxilla with ladnia. and galea comhined small tubercle on margin above lateral into one mesal clement; prcgula large to spine; Australia; Eucalyptus spp.; 5.5-6.3 very large, cleft between pregula and oral ITlI11..•...... l\i()t()JJlizt!lJOus ~a fossa nonexistent to shallow, equal to less than one-fourth pregula length; (Platypodi- 12(10). Eye elongate, 1.5 or more times longer than dae, Platypodini) 18 wide, almost flat; mesepistenllUTl flattened to concavely excavated, its upper (and 8(7). Proeoxae widely separated, each coxa very sometimes anterior) margin armed by a fine, large, longer than tibia; mesonotnm flat or conspicuous carina (carina absent in some evenly, transversely arched, without a con­ Spathidicerus); anterior nmrgin ofmesocoxal spicuous median carina; scutellum rather eavity acntely carinate, carina curving large, broad; male abdominal sternum ,S cephalad and ending in margin of mesepi­ sllbvertical, usually concavely excavated; sternum; pronotum more slender, 1.4-4.0 (Diapodini) 9 times as long as wide; preeoxal piece on prosternmll acutely pointed 13 Procoxae contiguous, each coxa shorter than tibia; mesonotllm with a conspicuous, acute, Eye slIbcircular, little if any longer than median carina (except flat, without a carina, wide, hemispherical; mesepisternurn con­ in Tesserocerus, Tesserocmnulus), scutellum vex, never armed by a carina; anterior mar­ small, slender, pointed; (Tesserocerrini) 10 gin of mesocoxaJ cavity never continued cephalad as a carina; pronotum stouter, 9(8) Anterior face of antennal club with a small 1.0-1.3 times as long as wide; precoxaJ area or line smooth, shining, often weakly piece on prosternum obtusc!y pointed; P1'o- elevated; base of female pronotum (occa­ notum and e1ytra rarely reticulate 16 sionally also male) with a band of many myeetangia pores or grooves; scutellum smaller, narrower, often depressed; frons 13(12). Mesonotum almost flat, never marked by a more sparsely pubescent in both sexes; median carina; eye shorter, oval to sllbtrian­ newly emerged female Ilsually with dehis­ gular in outline, 1.2-2.0 times as long as cent mandibular appendages; Africa to Tai- wide, anterior margin entire; Neotropical wan and Australia; 1.8-5.0 mm . species 14 ...... Diapus Chapuis Mesonotum conspicuously armed by a Antennal club uniformly pubescent to base; strongly elevated, acute, median carina; eye hase of pronotum ornamented in median very large, reniform, its anterior margin area by few coarse. mycetangia pores, never broadly emarginate, often two or more with grooves; scutellum larger, broader, times longer than wide; Afi'iean and Indo- higher; female frons usually ornamented by Australian spccies 15 tufts of very long setae; female dehiscent mandibular appendages usually absent; 14(13). Small, exceedingly slender species, body at India to Philippines and New Guinea; lenst 8.0 limes as lon~ as wide; lateral mar­ 1.7-4.0 mm Genyocerus Motschulsky gin of pronotum acute, lateral margin of posterior one-third of prosternum acutc, a 10(8). Antennal funicle 2--3-segmented, club with deep, longitudinal, pleural groove between large procurved, glabrous, basal area notal and sternal margins; female scape extending at least one-half length of dub, remarkably flattened and broadly extended margins and apical area minutely, closely mesad, dorsad, and caudad, pedicel inserted pubescent; protibiae armed by lateral spine one-third scape length from base; Costa at tarsal insertion and one additional subapi­ Rica to Cayenne; 4.0--4.2 mill . cal spine or ruga; elytral declivity gradual, ...... Tesserocmnulus Schedl 268 GREAT BASIN NATUHALIST [Volume 53

Larger, stouter species, 3.0-4.5 times as for reception of femur, anterior margin of long us wide; pronotum narrowly to suba­ impressed area never continuously carinate cutely rounded on lateral margins, prostcr­ or armed by a row of small spines (one Dum never with lateral margin costate, coarse nodule present on mctepistcrnum in without a narrow, pleural groove between male of some large Crossotarsus), surface of these margins; female scape slender, with impressed area with at least some setae; pedicel attached nelll" its apex, a slender, protibia of male armed by about four or elongate extension in a few species; :Mexico more coarse, transverse rugae, female either to Argentina; 3.0-11.0 mm .. similar to male or sometimes mostly cov­ ...... Tesserocerus Saunders ered by small, confused granules and usual­ ly one or two weak rugae near tarsal inser­ 15(13). Mosepistcrnum flat and unarmed by a cari­ tion; if present on female pronotum, myce- na (larger species) or concavely excavated tangia pores numerous 19 and its margin armed by a carina (smaller species); pronotal constriction (in which Metasternum and metepisternum near protibia moves) shallow, its posterior por­ metacoxa impressed for reception of femur, tion gradual, not extended ventrad, pleural anterior margin of impressed area either impression mostly below pronotal margin; continuously carinate or armed by a series protibia armed by only two coarse, trans­ ofsmall spines (absent in occasional females verse rugae; Indonesia to New Guinea and and in American allies of Euplatypus longu­ Philippines; 4.0-12.0 mm . lus), surface of impressed area glabrous; ...... Spathidicerus Chapuis protibiae of males and females similarly armed by rugae; spines on one or more Mesepisternurn always concavely excavat­ abdominal sterna (couplet 27) a common ed, its margin armed by a carina; pronotal feature; mycctangia pores variahle 25 constriction much deeper, its posterior por­ tion abrupt, with notum extending more 19(18). Male and female protibiae similarly armed ventrad; protibia armed by three coarse, by rows oftransverse rugae 20 transverse rugae; Afi'iea; 2.2-5.0 mm ...... Periommatus Chapuis :\Ilale protibia armed by transverse rugae, female protibia largely granulate, with no 16(12). Posterior one-third of pronotum with a more than one or two weak rugae ncar apex transverse band of numerous, small, closely ...... 23 placed mycctangia pores; anterior face of metatibia armed by only one transverse 20(19). Suture at apex of male clytral declivity ruga; striae more distinctly impressed, entire, deelivity variously convex, with or punctures clearly visible; female frons without armature of tubercles and spines; if impressed from eye to eye, central two­ present, female mycctangia pores on prono­ thirds of impressed area abruptly, deeply tum numerous; worldwide in most tropical excavated on a circular area; e1ytral declivi­ and subtropical areas, only 1 species in ty convex, rather steep, spines short, rather America; 2.5-10.5 mm Platypus Herbst inconspicuous; larger, stouter species; Afri- ca; 3.9-7.0 mm Chaetastus Nunberg Male declivity abruptly truncate, its margin Pronotal punctures uniform throughout, obtuse to very acutely costate on almost a mycctangia pores not discernible; anterior complete circle, apex sometimes strongly, face of metatibia armed by three or more attenuately narrowed, declivital face usually transverse rugae; striae weakly if at all concave; mycetangia pores variable 21 impressed, punctures evident or not; female frons variollsly impressed from eye to eye, 21(20). Elytral apex of male moderately to exceed­ without an abrupt, deep, central excavation; ingly attenuate, strongly narrowed to true smaller, more slender species 17 base of declivity, dehiscence of suture sometimes small, obscure, basal margin of 17(16). Male elytral declivity usually convex, very declivity usually more gradual, sometimes steep, usually not excavated, spines smaller, rounded; India and Australia to Japan and if evident; base of male declivity usually not NW North America; 2.4-6.0 mm " .. armed by spines; Central and South Ameri- ...... Treptoplatypus Schecll ca; 2.5--4.2 mm Cenocephalw, Chapuis Male elytral declivity much more broadly Male elytral declivity usually obliquely truncate, declivital base almost as wide as truncate and variously excavated, spines base of elytra, basal margin abrupt, obtusely much larger; base of male declivity usually to very acutely margined 22 armed by spines; Madagascar; 3.7-4.0 mm ...... Mitosoma Chapuis 22(21). Male sutural apex ofdeclivity usually entire, slightly dehiscent in one species; male e1y­ 18(7). Metasternum and metepisternum near tra not distinctly constricted before declivi­ metacoxa usually weakly or not impressed ty, costa at base of declivity obtuse to suba- 1993] REVISiON OF PLATYPODIDAE 269

cute, interstrial rows sometimes indicated 27(26). Visible male ahdominal sternum 5 armed by on upper porlion, at least a few setae pres­ a pair of widely separated spines; male ely­ ent, declivital face largely dull in most tral declivity shorter, steeper, its ventrolat­ species, shining in one; Malaya to New eral angles poorly developed and projecting Guinea; 2.8-4.5 mm Peroplatypus Wood little if any; male interstriae on posterior half of disc usually carinate; pronotlll1l Male sutural apex modestly to very strongly, never with l1lycetangia pores in either sex; very broadly emarginate; margin at base of small species, 1.9-3.5 mm; Mexico to Ar- male declivity moderately to strongly acute, gentina P1at!lscapulu8 Schedl face of declivity smooth, shining, glabrous, striae and interstriac never indicated; male Visible malc abdominal sternum 3 or 4 declivity with a distinct constriction slightly armed by a pair of widely separated spines; anterior to declivital base; India and Japan male elytral declivity more gradual, ventro­ to Australia and Micronesia; 2.8-5.5 mm . lateral angles more strongly produced; dis­ ...... Dinoplatypus Wood cal interstriae in male never carinate; pro­ notum on basal half often with a pair of my- 23(19). Male declivity very short to absent, usually cetangia pores in fcmale or in both sexes 28 subvertical, a row of spines usually arms base of declivity, venter of abdomen rising 28(27). Visible male sternum 3 armed by a pair of abruptly to meet elytra; male metepister­ spines; male declivity often steeper, shorter; num oflarger species often armed ncar pos­ mycetangia pores on pronotuIn often pres­ terior end by one rounded nodule; India ent in female or in both sexes; SE USA to and Australia to Taiwan and Hawaiian Venezuela; 2.0-5.5 mm Myop1at!lPU8 Wood Islands; 3.6-10.5 mm ...... Cmssotarsu8 Chapuis Visible male sternum 4 armed by a pair of spines; male pronotulH without mycetangia Male elytra strongly, more gradually pores, female with 1 pair of unusually large dec1ivous, venter of abdomen more nearly pores; (Juercus spp.; S USA to Chihuahua horizontal on segments 2-5; metepisternum and Nyarit in Mexico; 3.5-4.5 mm . never armed by a nodule 2,4 ...... Oxoplatypus 'Vood 24(23). Ventrolateral margin of male elytral declivi­ 29(26). Male clytra rather strongly dcclivous on ty evenly rounded, never serrate or dentate, posterior onc-third, declivity variollsly con­ its basal margin weakly armed, never den­ vex or obliquely impressed, with or without tate; male declivity usually convex, surface armature; venter of male abdomen rising dull; female pronotum never with myeetan- gia pores; Africa; 4.8-9.5 mm .. only slightly to mect apex ofclytra 30 ...... Trachyostu8 Schedl Male elytra descending little jf any before Ventrolateral margin of male declivity vari­ apex, declivity short, subvertical, if evident; ously serrate, dentate, or emarginate, its venter of male abdomen rising more than basal margin variously carinate or armed by one-halfdistance to meet apex 32 spines; male dec1ivitaI surface suheoneavely excavated; female pronotum with a pair of 30(29). Venter of male abdomen horizontal to ster­ myeetangia pores near median line on basal num 5, sternum 5 moderately to strongly half; S Mexico to Brazil; 5.D-7.5 mm . inflated, its apical one-fourth ascending ...... l\T~()trllcJ!!I()stus JBlrowlle rather abruptly to meet apex of elytra; male elytral declivity strongly convex, steep, 25(18). Anterior margin of impression on metaster­ unarmed or with small dentides on inter­ num and metepisternum for reception of striae 3, 7, 9, none on apical margin; myce­ femur armed by a series of small, pointed tangia pores on pronotum never present in spines (sometimes obscure or absent in either sex; Costa Rica to Brazil; 2.3-4.0 mm female Euplatypus); American or Madagas- ...... Platyphysus Wood car species, four from Africa 26 Venter of male ahdomen rising almost one­ Anterior margin of impression on metaster­ half distance to meet apex of c1ytra, declivi­ num and metepisternum armed by a com­ ty descending moderately, often variously plete or interrupted costa, rarely reduced to impressed and armed by spines 3 ] one (somewhat pointed) subcostate spinc; African species 34 31(29). Male declivity with ventrolateral angles usually formed and modestly produced, 26(25). Male visible abdominal sternum 3, 4, or 5 their apices never exceeding apical margin armed by a pair ofwidely (transversely) sep- at suture, margin between ventrolateral arated coarse spines 2 7 angles frequently armed by one or more pairs of dcnticlcs or serrations; mycetangia Male abdominal sterna 3-5 never armed by pores on pronotllm uncommon in female, spines 29 rare in male, when present, consisting of 270 GREAT BASIN NATURALIST [Volume 53

one pair or paired small dusters; Mexico to Male abdomen broadly concave, both trans­ Argentina; 2.3-10.0 mm...... Megaplatypus Wood versely and longitudinally, from base of sternum 1 to apex of 5, impressed area often Male declivity with ventrolateral angles elaborately pubescent; declivity descending more strongly produced, llsually exceeding very slightly, its margin armed by spines, apical margin at suture, projecting process interstriae 1 near its apex diverging laterad usually more slender and often with its apex moderately and descending slightly before 1i- or tridently armed, never with serrations its apex; female frons with a pair of small to or denticles on apical margin between moderately large concavities in lateral areas processes; pronotum often with one pair of between base of mandibles and antennal mycetangia pores in female, less common in insertions; Africa; 3.0-4.0 mm ., . male; mostly Mexico to Argentina, a few in ...... Triozastus Sehedl Africa and Madagascar, parallelu8 circum- tropical; 2.3-7.0 mm Euplatypus Wood 36(34). Male abdominal sternum 2 often armed by a pair of coarse, blunt spines, 3-5 ascending 32(29). Male declivity not descending, unarmed; and sometimes armed on one 01' more of male abdomimJ sternum 5 concave; small, these segments by small spines; male reticulate, very slender species, 5.0 or more declivity descending moderately, armature times as long as wide, upper surfaces usual~ rather inconspicuous; female frons concave- ly reticulate; numerous mycetangia pores on ly impressed; Africa; 3.5-4.5 mm .. pronotum, if present; sexual dimorphism ...... lldr~l>Cl1?lllttllJ~ StJrohllleyeI' obscurc; Australia to Malaya; 1.7-2.4 mm...... Baiocis Browne Male abdominal sternum 2 transversely car­ inate, carina moderately to extremely, Less slender species; sexual dimorphism strongly elevated and either continuous or conspicuous, male declivity always with interrupted near median line; declivity usu­ small spines; when present, mycetangia ally armed on its basal margin by dorsoven­ pores limited to one pair; American species... 33 trally flattened costae, these costae inter­ rupted at strial intervals, declivity below 33(32). Male declivity with two pairs of serrations these spines weak to nonexistent; female on ventrolateral margin, these serrations frons variously sculptured, often elaborately usually connected by a carina, median pair ornamented by setae; Africa; 2.5-7.0 mm ..... (often both) on apical margin; one pair of ...... Dolwpygus Sehedl myc.:etangia pores on pronotmn often pres- ent; Costa Him to Brazil; 2.8-4.5 mm .. COPTONOTINAE ...... Epiplatypus Wood The classification of Coptonotinae remains Male declivity with only one pair of serra­ tions on ventrolateral margin, a carina as presented in Wood in Wood & Bright extending dorsad from this spine to a spine (cl992), containing the following: Coptonotini on interstriae 3 at base of declivity, basal (CoptonatWi Chapuis, 2 Neotropical species; margin at apex of disc usually armed by Protohylastes Wood, 2 Australian species; smull spines on interstriae 1, 3, 5; mycetan­ Scalytatarsus Schedl, 1 African and 1 Aus­ gia pores on pronotum never present; S Mexico to Argentina; 2.2-4.2 mm .. tralian species); Mecopelmini (Mecapelmus ...... J'~l~~~~ 1'1,",~ Blackman, 1 species from Panama; Proloplaty­ pWi Wood, 1 species from New Guinea); and 34(25). Male abdomen with sternum 2 normal, ster­ Schedlarini (SchedlariWi Wood, 1 species from na gradually ascending from 1-5, unarmed; Mexico). Mecopelmus zeteki Blackman is declivity descending slightly to moderately; female frons often variously concave 35 known only from specimens collected within 2 km ofthe Panama CanaL It is quite probable Male abdomen with sternum 2 abnonnally that this species was introduced from another long, armed or abruptly angled on 2, part of the world, possibly New Guinea where ascending from 3-5; female frons often the only known relative occurs. densely pubescent 36

35(34). Male abdomen with sterna 1-5 transversely TESSEROCEiuNAE convcx, with normal setation; elytral decliVi­ ty convex, descending about one-half dis­ The Tesserocerinae are divided into two tance to meet ascending abdomen; decIivital trihes as presented in Wood in Wood & Bright tubercles small, inconspicuous; female frons (cl992): Diapodini (Diapus Chapuis, 39 broadly and shallowly to strongly concave; Africa, Madagascar; 2.5-5.5 mm .. species from India to Australia; Genyocerus ...... •...... <::!lli~,.olJ(JllJ~ 1;tI'ohllle"el' Motschulsky [= DiacavWi Schedl], 24 species 1993] REVISION OF PLATYPODIDAE 271 from India and Sri Lanka to Philippines and (cl992). Foremost among these is the aban­ New Guinea); and Tesserocerini. donment of the genus "sektionen" of Chapuis A divided maxilla into separate lacinia and (1865), Strohmeyer (1912, 1914b), and Scbedl galea lobes occurs in the primitive Cur­ (1972). This non-Linnaean category was culionoidea (Anthribidae, Nemonychidae) apparently below the rank of subgenus but having two gular sutures (Crowson 1955, above the rank of species and was used liber­ 1968, Wood 1986). Among the higher Cur­ ally by Schedl with little objectivity. These culionoidea, those with only one gular suture, "sektionen" are here replaced by a new classi­ divided lacinia and galea lobes occur only in fication ofgenera. parts of Attelabidae, Rhynchitidae, and Platy­ The Platypodinae, as presented here, podidae (Tesserocerinae; Wood 1986:8, fig. 8). appear to represent a recent, active, evolu­ In all tbree ofthese families the taxon contain­ tionary explosion in which sharply delineated ing all species witb separate lacinia and galea generic groups do not exist. For this reason all is given subfamily status. Strohmeyer (1912, are placed in one tribe, Platypodini. Schedl's 1914b) appreciated this fact and recognized (1972:83) attempt to characterize his Crosso­ the subfamily Tesserocerinae. Schedl (1972) tarsini as distinct from his Platypodinae was was not a student of evolution and did not based on a character (sexual dimorphism of acknowledge the existence of this character in the protibiae) that did not occur throughout Platypodidae. the group he attempted to characterize, nor To the Tesserocerini of Strohmeyer (1912, was it limited to his Crossotarsini. Another set 1914b) two genera are added here, Platytarsu­ of characters was needed to divide his Platy­ Ius Schedl and Notaplatypus Lea, on the basis podinae. of the deep cleft between the pregula and the On the posterior portions of the metaster­ margin of the oral fossa (specimens for dissec­ num and metepisternum of some Platypodini tion of the maxilla were not available). The is a feeble to very strong, often glabrous Tesserocerini now contain (Wood in Wood & impression for the reception of the metafe­ Brigbt cl992) the following: Platytarsulus mur. The anterior and lateral margins of this Schedl (8 species from Malaya and Borneo); impression may be armed by (1) a continuous Notoplatypus Lea (1 species from Australia); carina (African species) or (2) a series of Tesserocranulus Schedl (1 species from Costa minute spines (American species, with a few Rica to Cayenne); Tesserocerus Saunders eastern hemisphere exceptions). Those Platy­ (= Damicerus Spinola, Tesseroplatypus podini that lack this impression and its carina Schedl, Tesserocephalus Schedl) (30 species or spines also share other features generally from southern Mexico to Argentina); not found in the other group. It should be Spathidicerus Chapuis (7 species from Suma­ mentioned that occasional females (American tra to Pbilippines and New Guinea); Periom­ species) and about a dozen species allied to matus Chapuis (=Asetus Nunberg, Setanus Euplatypus longulus (Chapuis) (all are Ameri­ Nunberg) (52 species from tropical Africa); can species) lack the impression and spines Chaetastus Nunberg (=Symmerus Chapuis) (7 even though they (otherwise) clearly belong to species from tropical Africa); Cenocephalus the generic group with the impression and Chapuis (13 species from southern Mexico spines. Conversely, several of the largest and Hispanola to Brazil); and Mitosoma Cha­ species of Crossotarsus have one small, round­ puis (=Platypicerus Nunberg, Coecephalonus ed nodule on the male metepisternum Schedl) (26 species from Madagascar). although they clearly belong to the generic Schedl (1972) did not recognize the Tesse­ group without the impression or spines. rocerinae as a subfamily, but fragmented the Among those groups treated here as genera group into his Diaporinae [sic], Periommati­ that lack the impression and its armature, all nae, and Platypodinae. (mostly females) that have mycetangia pores on the pronotum have numerous pores. PLATYPODINAE Among those genera with the impression and spinelike annature, most of those species (pri­ Introduced here are radical changes in the marily females) with mycetangia pores on the classification of Platypodinae tbat were found pronotum have only one pair, although a few too late for inclusion in Wood & Bright large Megaplatypus have several, and the few 272 GREAT BASIN NATURALIST [Volume 53

Baiacis with pores have many. All of those lreptoplatypus Schedl species with mycetangia pores on the prono­ The genus Treptoplatypus Schedl was tum and also with a carina on the metaster­ based on CrOs8otarsus trepanatus Chapuis. num-metcpisternum impression (African Scbedl (1972:245) also included circulicauda species) have many pores. Browne, fischeri Strohmeyer, multiparus Scbedl, qoodriporu~ Schedl, and subaplanatus Platypu~ Herbst Schedl, all (five) ofwhich are nnknown to me. As indicated above, Treptoplatypus taxicornus Tbe genus Platypus Herbst (= Cylindra (Schedl) is here transferred back to Platypus. Illiger, Stenoplatypus Strohmeyer, Plntypinus DESCRIPTION.-A member of the Platypo­ Schedl) as defined bere is greatly reduced in dini near Platypus, Treptoplatypus is distin­ the number of included species from that list M guished by the strongly narrowed male elytral ed by Schedl (1972:169-242) and Wood & declivity that is rather abruptly, obliquely Bright (cl992). To these synonyms is addcd truncate and dehiscent at the sutural apex. Austroplatypus Browne (1971:49), ncw syn­ The male elytral apex is usually strongly onymy. It also appears that Dendroplatypus attenuate, and the male declivity is usually Browne (1955:365) bclongs herc (only females concave. Mycetangia pores on the female were available for study). Neotrachyostus pronotum are numerous. quadrilobus (Blandford) is here transferred CONTENTS.-In addition to trepanatus, I from Neotrachyostus back to Platypus. Platy­ here transf"r from Platypus to Treptoplatypus pus taxicorm:s Schedl belongs here, not in the species abietis (Wood), australis (Cbapuis), Treptoplatypus where it was placed by Schedl biflexuosus (Schedl), micuras (Schedl), solidus (1972:245). (Walker), and wilsoni (Swaine). It is probable DESCHIPTION.-Platypus Herbst is a mem­ that some (not all) species placed by Schedl ber of the Platypodini, as defined in the above (1972:197-199) in Platypi oxyuri sbonld also key to genera, in which the posterior portions be transferred here, as well as longipennis of the metasternum and metepisternum are Montrouzier (Schedl 1972:196). Additional not impressed or armed (key couplet 18a) and studies are needed to determine exactly the protibiae are not sexually dimorphic (key which species should and should oot bc couplct 19a). The male sutural apex on the added to this genus. elytral declivity is not dehisccnt. Mycctangia DISTRIBUTION.-India and Japan to Aus­ pores when present on the pronotum (mostly tralia and NW North America. females) are numerous. Peroplatypus, n. g. CONTENTs.-Included here in this group are the following "sektionen" of Platypus as DIAGNOSIS.-This genus is a member of listed by Schedl (1972:169-242): Platypi api­ the Platypodini near Treptoplntypus, but it is cali (1 sp., Fiji), Platypi gerninati (3 spp., New distinguishcd from tbat genus hy the broad Guinea), Platypi hirtelli (22 spp., India to Aus­ elytral declivity that is obliqnely truncate, with the suture entire (slightly dehiscent in tralia and Philippines), Platypi lunati (15 spp., one species). It is distinguisbed from Dino­ india to Australia), Platypi rnesoadjuncti (3 platypus by the absence ofan elytral constric­ spp., Malaya to New Guinea), Platypi tion immediately cephalad from the declivity, paraspinulosi (5 spp., Africa), Platypi pseu­ and hy the presence of setae on tbe face of the dospinulosi (12 spp., Malaya and China to male declivity. New Guinea), Platypi punetati (2 spp., lndia DESCRIPTION.-Metasternum and metepi­ to New Guinea), Platypi semiopaci (9 spp., sternum without an impression or armature Australia to New Guinea), Platypi spinulosi for reception of the metatibia. Tbe malc cly­ (13 spp., Africa), Platypi suleali (60 spp., tral declivity is broadly, obliquely truncate, Europe, India, and Japan to Australia). This not preceded by a transverse constriction; the reduces the 808 species of Platypus listed in suture is entire (one slight exception); the Wood & Bright (cl992) to 121 species. declivital face is ornamented by setae (either DISTlHBUTION.-Europe and Africa to bairlike or scalelike); the costa at the base of Japan and Australia, 1 species (quadrilobus the male declivity is obtuse to subacute, and Blandford) of dubious affinity in Costa Rica. the interstrial rows are sometimes indicated 1993] REVISION OF PLATYPODIDAE 273 on the upper portion. The male declivitaJ f~lce declivity is moderately reduced to almost is usually dull (shining in one species). ahsent (a row of dorsoventrally flattened CONTENTs.-Type-species: Platypus trun­ spines arms its basal margin); the abdomen catipennis Schedl. Included here are the ascends rather strongly to meet the apex. The Platypi sulcato-truncati (5 spp., Borneo, New males of several of the larger species have a Guinea) and Platypi truncatipenni (6 spp., rounded nodule on the metepisternum. The Borneo, Sumatra, New Guinea) of Schedl female pronotum has numerous mycetangia (1972:211-212). Of these, only platypoides pores. (Browne), truncaticauda (Schedl), truncatigra­ CONTECoITs.-lncluded here are the follow­ nasus (Schedl), and truncatipennis (Schedl) ing groups as listed by Schedl (1972:96-112): were at hand for study. Grossotarsi alternante-depressi (1 sp., Philip­ DISTRIBL'TION.-Malaya to New Guinea. pines), Crossotarsi angulati (4 spp., India, Japan, Ncw Guinea), Crossotarsi harbati (11 Dinoplatypus, n. g. spp., Malaya to Philippines and Australia), DIAGKOSIS.-The genus Dinoplatypus is Crossotarsi coleoptrati (12 spp., India to Japan distinguished from Peroplatypus \\Tood, above, and New Guinea), Crossotarsi subdepressi (20 by the suhvertical, obliquely truncate male spp., India to Taiwan and Australia), Cros8o­ clytral declivity with the sutural apex modest­ tarsi genuini (20 spp., India to Philippines and ly to very strongly, very broadly emarginate, Australia), Crossotarsi nitiduli (4 spp., Malaya and with the subvertical face moderately to to New Guinea), Crossotarsi ventriC0111.i (14 strongly concave, brightly shining, and with­ spp.. India to Japan and New Guinea), out punctures or setae; the upper margin of Crossotarsi incertae sedis (.3 spp., Java to the male declivital face is usually acute, and Philippines, 1 sp. of doubtful affinity in there is a distinct constriction immediately Africa). cephalad from its base. DISTHlBUTION.-India to Japan and Aus­ DESCHIPTlON.-The male clytral declivity tralia, externedentatus has extended its range is subvertically truncate; its upper margin is through modern commerce to Hawaii and has acute; its face is broadly, subcircularly con­ been intercepted in additional areas. cave; its surface is brightly shining, impunc­ Trachyostus Schedl tate, glabrous, with a substantial, often elaho­ rate, emargination at the sutural apex. The This genus is allied to Crossotarsus, but it male declivity has a distinct, transverse con­ is confined to Africa and Madagascar. striction immediately cephalad from its base. DESCRIPTION.-Trachyostus is allied to The female pronotum has numerous mycetan­ Crossotarsus as indicated by the similarly sex­ gia pores. ually dimorphic protibiae. The male elytral CONTEl\TS.-Type-species: Platypus cupu­ declivity is usually convex (rarely flattened), latus Chapuis. Included hcre are thc Platypi evenly rounded, never serrate or dentate, and cupulati (29 spp.) of Schedl (1972:208-211). the surface is usually dull. The venter of the DISTRIBUTION.-India and Japan to Aus­ abdomen ascends little, ifany, to meet the ely­ tralia and Micronesia. tral apex. Mycctangia pores are never present on the pronotllm. Crossotarsus Chapuis CONTENTS.-Included here are the 13 The genus Crossotarsus Chapuis, as treat­ species from tropical Africa and Madagascar ed here, is essentially as listed in Schedl that were listed by Schedl (J.972:88-89) and (1972:96-112) and Wood & Bright (c1992), Wood & Bright (cl992). although it may become necessary to add to it OrSTlUBUTlON.-Tropical Africa and Mada­ all or part of Carchesiopygus Schedl (not seen) gascar. and Crossotarsinulus Schedl (not seen). Neotrachyostus Browne DESCHIVrIO:\l.-Crossotarsus is a member of the Platypodini, near Platypus, except that The genus Neotrachyostus Browne, as used (key couplet 19) the protibiae are sexually here, is essentially as listed in Schedl dimorphic (male "vith the usual transverse (1972:90-92) and Wood & Bright (cl992) rugae, female with most of the basal fugae except that Platypus qnadrilobus Blandford is replaced hy confused granules). The male here transferred back to Platypus. 274 GREAT BASIN NATURAUST [Volume 53

DESCRIPTIO .-The sexually dimorphic usually has one pair of mycetangia pores in protibiae of Neotrachyostus suggest a close the female; they are sometimes present in the relationship to Trachyostus. The male e1ytral male. declivital surface is never dull; it is variously CONTENTS.-Type-species: Bostrichu" flav­ impressed or excavated, with the ventrolateral icomis Nlbricius. Included here are flapi<;or­ margin serrate, dentate, or emarginate; its nis (Fabricius) (S USA to Cuba) and Schedl's base is variously carinate or armed by spines. (1972:220) Platypi bilobati (5 spp., Mexico to The female pronotum has one pair of my­ Costa Rica). cetangia pores. DISTRIBUTION.-Southeastern USA and CONTENTs.-Schedl (1972:92) and Wood & Cuba to Mexico and Venezuela. Bright (cl992) list 14 species. Oxoplatypus, n. g. DISTRIBUTION.-Southern Mexico to Brazil. DIAGNOSIS.-This genus is a memher of the Platypodini near Platyscapulus. It is dis­ Platyscapulus Schedl tinguished from Platyscapulus by tlle pres­ The genus Platyscapulus Schedl (=Platy­ ence of a transverse pair of large spines that scapus Schedl 1939:397, 399, Costaroplatus arm male visible abdominal sternum 3, and by Nunberg 1963:109) contains a group ofAmer­ the absence ofspines on sternum 5. ican species formerly assigned to Platypus. DESCRIPTION.-This genus is established Platyscapt/lus is here removed from synonymy to contain one known species. It is a represen­ with Platypus and is given full geneiic rank. tative of the Platypodini with the metaster­ DESCRIPTION.-As defined here Platy­ num-metepisternum impression armed by scapulus contains those species formerly small spines on the anterior margin, and male assigned to Platypus that have the metaster­ visihle abdominal sternum 3 is armed by a num-metepisternum impression anned on its transverse pair of large spines. The female anterior margin by a series of small spines and pronotum bears one unusually large pair of also have a pair of spines that arm visible male mycetangia pores; the male pronotum is with­ abdominal sternum 5 (Schedl 1972:195, fig. out pores. 49). The male e1ytral declivity is usually short, CONTENTS.-Type-species: Scolytus quad­ steep, and has the ventrolateral angles rather ridentatus Olivier. One species is known. poorly developed, projecting little, ifany. The quadridentatus (Olivier) (=blanchardi Cha­ male elytral interstriae are usually carinate on puis, disciporus Chapuis). the posterior half of the disc. The pronotum DISTRIBUTION.-Southeastern USA to never has mycetangia pores in either sex. northern Mexico,"in Quercus spp. CONTENTs.-Included here are Sched!'s Platyphysus, n. g. (1972:235) Platypi coswUati (13 spp., S Mexico to Brazil), Platypi abckJminales (Schedl 1972: DIAGNOSIS.-This genus is a member of 195) (3 spp., Costa Rica to Guyana), and the Platypodini having the metasternum­ PlatlJPi neocosteUati (SchedlI972:195) (2 spp., metepistemum impression anned on its ante­ Venezuela and Guyana to Brazil). rior margin by small spines, but none of the DISTRIBUTION.-Southern Mexico to visible male ahdominal sterna is armed by Brazil. spines. Platyphysus is distinguished from allied genera by the strongly convex, steep Myoplatypus, n. g. male elytral declivity that is almost unarmed, DIAGNOSIS.-This genus is distinguished and by the horiwntal venter of the abdomen from the closely allied Oxyplatypus Wood, with visible sternum 5 inflated, its posterior below, by the occurrence of a pair of large one-fourth ascending rather abruptly to meet spines on male visible abdominal sternum 4, the apex ofthe elytra. and by the absence of spines on other sterna. DESGRIPTION.-In this genus visihle male DESCRIPTION.-This genus is a member of ahdominal sternum 5 is strongly inflated the Platypodini that have a metasternum­ (moderate in female), with its posterior one­ metepisternum impression armed by small fourth ascending to meet the apex of the ely­ spines and a transverse pair of spines on male ira. The elytral declivity is convex, steep, and visible ahdominal sternum 4. The pronotum descends further than in related genera; male 1993] REVISION OF PLATYPODIDAE 275 armature is sparse and rather small. The more strongly produced ventrolateral angles metasternum-metepisternum impression is of the male declivity that exceed the level of armed by small spines as in related genera. the sutural apex. CONTENTs.-Type-species: Platypus obtu­ DESCRIPTION.-This genus is a member of sus Chapuis. Also included here are Schedl's the Platypodini having the metasternum­ (1972:187) Platypi declivi (4 spp., Brazil) and metepisternum impression armed on its ante­ Platypus pauteriae Wood. rior margin by small spines. None of the visi­ DISTRIBUTION.-Costa Rica to Venezuela, ble abdominal sterna are armed by spines. in Pouteria spp. The male ventrolateral angles of thc declivity are extended caudad into a pair of processes Megaplatypu8, n. g. that exceed the sutural apex (apices of each of DIAG~OSIS,-This large group of American these processes arc usually bi- or tridentate, species, formerly placed in Platypus, is and never with serrations or denticles on the diverse and is distinguished with some diffi­ apical margin between these processes). The culty. From Euplatypus Wood, bclow, it is dis­ pronotum often has one pair of mycctangia tinguished by the more poorly formed and pores in the female or in both sexes. much less strongly produced posterolateral CONTENTs.-1ype-species: Bostrichus par­ angles of the male elytra (key couplet 31); one allelus Fabricius. Also included here are or two pairs of small denUdes sometimes arm Sched!', (1972:230-234) Platypi tlispinati (39 the apical margin between these angles. Myce­ spp., USA to Argentina, MadagascaI~ tropical tangia pores are uncommon (female) or rare Africa, Australia, Sri Lauka, etc.) and Sehed!'s (male) but may consist of one pair or a pair of (1972:205) Platypi caudati (19 spp., S Mexico clusters of pores (perhaps 4 to 12 on eaeh to Argentina). Some of the caudati group from side). tropical America lack the small spines that DESCRIPTION.-This is a genus of Platypo­ arm the metasternum-metepisternum impres­ dini having the metasternum-metepisternum sion in one or both sexes. impression armed by small spines; they lack DISTHIBUTlON.-Southern USA to Argenti­ spines on the visible male abdominal sterna. na, a few in Africa, Madagascar. Euplatypus The male declivity descends at least half the parallelus (Fabricius) has been carried distance to meet the abdomen, its lateral through modern commerce worldwide in angles are rather poorly produced (usually tropical areas (Wood & Bright cl992: they do not exceed the apex of the suture), 1664-1668). It has also been intercepted in and the apical margin betweeu these angles Australia and India in recent months. sometimes is armed by one or two pairs of Baiocis Browne small denticles. The pronotum usually is with­ The genus Baiocis Browne as treated here out myeetangia pores, but one pair or multiple is essentially as it was established by Browne pores are sometimes present (particularly in (1962:651) and listed by Wood & Bright the female). (cl992), except that Platypus kuntzeni Schedl CONTENTs.~Type-species: Platypus denta­ apparently belongs in Crossotarsus. tus Dalman. Also included here are Sched!'s DESCRIPTION.-This genus is a member of (1972:238-242) Platypi plicati (82 spp., S Mex­ the Platypodini having the mctasternum­ ico to Argentina), Sehed!'s (1972:186-189) metepisternum impression armed on its ante­ Platypi discoidales (4 spp., S Mexico to rior margin by small spines. The species are Brazil), Sehed!'s (1972:184) Platypi punctato­ small, usually reticulate, very slender, with sulcati (1 sp., Guatemala to Panama), Sehed!'s sexual dimorphism obscure. The male elytral (1972:229) Platypi pseudacaudati (1 spp., declivity is unarmed and it descends feebly, if Guyana to Brazil), Platypus nudatus Wood at ,JI. The visihle male abdominal stcrnum .5 (Colombia), 1'. pemudus Sehedl (Guyaua), and is concave. Mycetangia pores, when present 1'. simpli.cifarmis Wood (Costa Hiea). on the pronotum, are numerous. DISTRIBUTION.-Mexieo to Argentina. DISTRIBUTION.-Australia to Malaya. Euplatypus, n. g. Epiplatypus, n. g. DIAGNOSIS.-This genus is distinguished DIAGNOSIS.-This genus is a member of from Megaplatypus \Vood, above, hy the much the Platypodini having the metasternum- 276 GREAT BASIN NATURALIST [Volume 53 metepisternum impression armed by small the metasternum-metepisternum impression spines on its anterior margin. It is distin­ continuously costate. The abdomen ascends guished in the male from Megaplatypus gradually and moderately to meet the elytra. Wood, ahove, and Teloplatypus Wood, helow, The male visible abdominal sterna are trans­ hy the unique structure of the male elytral versely convex, sternum 2 is not enlarged or declivity. modified. The male elytral declivity is convex; DESCRIPTION.-This genus is allied to Telo­ moderately steep, and with tubercles small platypus hut is distinguished hy the presence and inconspicuous. The female frons is broad of two pairs of serrations on the ventrolateral and shallowly to moderately concave. margin of the male elytral declivity; these ser­ CONTENTs.-Wood & Bright (cl992) list 14 rations are usually connected by a carina; the speCIes. median pair (often hath pairs) is on the apical DISTRIBUTION.-Africa to Madagascar. margin. One pair of mycetangia pores is often present on the female pronotum or ~n both Triozastus Schedl sexes. The genus Triozastus Schedl, as treated CONTENTs.-Type-species: Platypus des­ here, is essentially as listed by Schedl ceplor Wood. Also included here are Platypus (1972:246-248) and Wood & Bright (cl992). annexus Wood, P. applanatus Wood, P. There appears to be considerable confusion in deplanatus Wood, P. eugestus Wood, P. eximius this genus on how to interpret individual and Wood, P. filaris Wood, P. jamacensis Bright, P. populational variability into taxonomic cate­ secus Wood, P. spectus Wood, P. vegestus gories. Wood, and apparently most of Schedl's DESCRIPTION.-This genus is distinguished (1972:213-214) Platypi complanati. from Cylindropalpus Strohmeyer by the male DISTRIBUTION.-Costa Rica to Brazil. abdomen being broadly concave (hoth trans­ Teloplatypus, n. g. versely and longitudinally) from the base of visible sternum 1 to the apex of 5, this con­ DIAGNOSIS.-This genus is distinguished cave area being often elaborately pubescent. from Epiplatypus Wood, ahove, hy the unique The male elytral declivity descends only structure of the male elytral declivity as slightly, and its basal margin is armed by defined in the above key to genera. spines; interstriae 1 near its apex diverges lat­ DESCIUPTION.-This genus is a member of erad moderately then descends slightly before the Platypodini having the metasternum­ its apex. The female frons bears a pair of small metepisternum impression armed on its ante­ rior margin by small spines. The male elytral to rather large concavities in the lateral areas declivity has only one pair of serrations on the between tbe bases of the mandibles and the ventrolateral margin, with a carina extending antennal insertions. dorsad from this spine to a spine on interstriae CONTENTs.-Wood & Bright (cI992) list 7 3 located at the base of the declivity; the speCIes. declivity descends only slightly, and its basal DISTRIBUTION.-Tropical Africa. margin is usually armed by small spines on Mesoplatypus Strohmeyer interstriae 1, 3, and 5. Mycetangia pores are never present on the pronotum in either sex. As treated here, the genus Mesoplatypus CONTENTs.-Type-species: Platypus con­ Strohmeyer is based on Wood & Bright cinnus Blandford. Included here is Schedl's (cl992) and on Schedl (1972:165-168). (1972:218-219) Platypi terminati (16 spp.). DESCRIPTION.-This genus is a member of DISTRIRUTION.-Southern Mexico to that portion of the Platypodini having a Argentina. costate anterior margin of the metasternum­ metepisternum impression and having visible Cylindopalpus Strohmeyer male abdominal sterna 2, 3, or 4 armed by The genus Cylindropalpus Strohmeyer, as spines. In some members male sternum 2 treated here, is essentially as listed by Browne bears at least a partial b"ansverse carina that is (1962:650, 655), Schedl (1972:131-134), and reminiscent of Doliopygus. The female frons Wood & Bright (cl992). is concavely impressed (in all species?). DESClUPTION.-This genus is a member of CONTENTs.-Wood & Bright (cl992) list 17 the Platypodini having the anterior margin of speCIes. 1993J REVISION OF PLATYPODIDAE 277

DISTRIBUTION.-Tropical Africa. · 1968. A natural classification of the families of --C::-QI~eoptera.Edition 2. E. W. Classey Ltd., Hampton, Doliopygus Schedl England. 195 pp., 213 figs. FABRICIUS, J. C. 1792. Entomologica systematica emenda­ The genus DoliopygW! Schedl (=Scutopy­ ta et aucta, secondum classes, ordines, genera, gus Nunberg, PygodoliW! Nunberg, Mixopy­ species adjectis synonymis, lods, ohservationihus, gW! Nunberg, MesopygW! Nunberg), as treated descriptionihus. Praff:, Hafniae, vol. 1, pt. 2. 538 pp. here, is essentially as listed by Schedl HERBST, J. F. W. 1793. Natursystem alIer bekannten in­ (1972: 143-164) and by Wood & Bright und auslandischen Insekten, als eine fortsetzung der (cl992). von Buffonschen Naturgeschichte. Der Kafer, vol. 5. DESCR[PTION.~This 392 pp., 16 pis. genus is allied to HOPKINS, A. D. 1909. Contributions toward a monograph MesoplatypW! Strobmeyer but is sharply dis­ of the scolytid beetles. 1. The genus Dendroctonw. tinguished by characters of the male U.S. Department of Agriculture, Bureau of Ento­ abdomen. Male visible abdominal sternum 2 mology, Technical Bulletin 17(1). 164 pp., 8 pIs., 95 has a strongly developed, transverse carina figs. that is sometimes divided at the median line. __:;--,' 1911. Contributions toward a monograph of The sternum caudad from this carina ascends the bark weevils ofthe genus pwsodes. U.S. Depart­ ment of Agriculture, Bureau of Entomology, Tech­ abruptly in union with sterna 3, 4, and 5 to nical Bulletin 20(1). 68 pp., 22 pis., 9 figs. form a subvertical, strongly concave, subcir­ LATREILLE, P. A. 1807. Genera Cfustaceorum et insecto­ cular face that functions in the removal of rum secundum ordinem naturalem in familias dis­ frass from the gallery entrance hole. The male posita, inconibus exemplisque plurimius explicata. declivity is reduced to obsolete; its basal mar­ Paris. Vol. 2. 280 pp. ROBERTS, H. 1993. Diapodini of Papua New Guinea gin is armed by a row of dorsoventrally flat­ (Platypodidae). Bishop Museum Occasional Papers tened costae (derived from spines) that are 35:1-39. interrupted at the strial intervals. The female SCHEDL, K. E. 1939. Die Einteilung und geographische frons is variously sculptured and may be elab­ Verbreitung der Platypodidae. International Con­ orately ornamented by setae in some species. gress of Entomology, Proceedings 7(1):377-410. Mycetangia pores on the pronotum are __~:-:' 1962. Fam. Coptonotidae. Coleoptera. Genera absent. Insectorum de P. Wytsman (Mercurius, Anvers), Fascicle 215. 13 pp., 1 pl. CONTENTS.-Wood & Bright (cl992) list __::-;-' 1972. Monographie der Familie PJatypodidae, 142 species. Coleoptera. W. Junk, Den Haag. v + 322 pp. DISTRIBUTION.-Tropical Africa. SHUCKARD, W. E. 1840. The British Coleoptera delineat­ ed by W. J. SplY. 83 pp., 94 figs. Reprinted in 1861. STROHMEYER, H. 1912. Familia Platypodidae. Pars LITERATURE CITED 44:1-26 in W. Junk and S. Schenkling, Coleoptero­ rum Catalogus. Berlin. BLACKMAN, M. W. 1944. A new genus and species of ___. 1914a. Coleoptera: Fam. Chapuisidae. P. Wyts­ Coleoptera from Panama. Entomological Society of man, Genera Insectorum, Bruxelles, Fasc. 162. 6 Washington, Proceedings 46(3):76-80, pI. 7, figs. pp., 1 pI. 1-5. · 1914b. Coleoptera: Fam. Platypodidae. P. BROWNE, F. G. 1955. Synonymy and descriptions ofsome --"W"y-"tsman, Genera Insectorum, BruxelIes, Fasc. 163. oriental Scolytidae and Platypodidae (Coleoptera). 55 pp., 12 pIs. Sarawak Museum Journal 6:343-373. WOOD, S. L. 1957. A new generic name for and some bio­ ____. 1962. Taxonomic notes on Platypodidae (Cole­ logical data on an unusual Central American beetle optera). Annals and Magazine of Natural History, (Coleoptera: Platypodidae). Great Basin Naturalist scr. 13,4:641-656 (1961). 17:103-104. __~~. 1971. Austroplatypus, a new genus of Platypo­ · 1966. New records and species of Neotropical didae (Coleoptera), infesting living Eucalyptus trees --"'p'-laC'typodidae (Coleoptera). Great Basin Naturalist in Australia. Commonwealth Forestry Review 26,45-70. 50:49-50. · 1973. On the taxonomic status of Platypodidae ____. 1972. Larvae of the principal Old World gen­ -~-an-d'- Scolytidae (Coleoptera). Great Basin Naturalist era of Platypodinae (Coleoptera: Platypodidae). 33,77-90. Royal Entomological Society of London, Transac­ · 1982. The bark and ambrosia beetles of North tions 124:167-190. ---,n-d'Central America (Coleoptera: Scolytidae), a tax­ CHAPUIS, F. 1865. Monographie des Platypodides. Des­ onomic monograph. Great Basin Naturalist Memoirs sain, Liege. 344 pp. 6. 1359 pp. CROWSON, R. A, 1955. The natural classification of the _~=_. 1986. A reclassification of the genera of Scoly­ families of Coleoptera. Nathaniel Lloyd & Co., Lon­ tidae (Coleoptera). Great Basin Naturalist Memoirs don. 187 pp. 10.126 pp. 278 GREAT BASIN NATUllALiST [Volume 53

WOOD, S. L., ANU D. K RRICIIT. In. cL992. A catalog or lunifer Schedl Scolytidac and Plalypoclidae (Coleoptera), pMt 2: luzoniew: Schedl taxonomic index. Creat Basin Naturalist Memoirs mirlUtissimus SchcdI 13:1084-1240. Vol. 2. rCopyrightcd 13 Det:crnber m,jobergi Scheel! 1992 by Brigham Young University]. mod.e,stus Rlandford morigerus Schedl rmJttr(Wu.~ School neoplicatus Schedl PAIlTJAL CIIECKLIST OF PLATYPODlNAE niiftrtUIi Murayama obtusi{)ennis Schedl As an aid to the interpretation of the above omissus Schedl changes, the following list of valid names in opllCideclivis Scheell Platypodinae is presented. Only valid generic opocifrons SchcdO sttbplicatus Sehcdl jamoni Chu\1uis subsccretus Browne juvencus SchecU subsidalius Schedl kalshoveni Sched! subsimilis Sehedl kitlShuensl.~ Mllrayama suffOdiellS Sampson kWpperi<;hi (Sol1"dl) tasnul.rl.ir.us Schedl 1.fltedP..clivis Schedl taricomis Sclledl line_tins Schc

unifurmis School Crossotarsus uubUI. (School) (Sec Wood & Bri~lt cl992:1195-12(9) uerelunatus (Beeson) veR<:Ulus Schcdl CarcJU'Mopygw; oedu. Strohmeyer (S~ Wood & Bright cl992:1209-1210 vetulus School webberi Schcdl Cro.wwtarsinulus westwoodi Chapuis (SL,,{~ Woud & Rl"ight cl992: 121.0)

Treptup1at!IPus Trachyustus ahie'l. (Wood) (See Wood & Bright d9Q2.:121O-1213) australis (Chapuis) biflexuosus (Schedl) Neotr-achyostus circulicauda Browne (Sec Wood & B';ght c19n1213-1214) flScheri {Stmhmeycr) micn~ (Schedl) Platysc.apulus nwltiporu, SchooL ahclitlJn, (Wood) ","ul,.;poru, Schedl ,r (Chapuis) plat1JPOides (Brown.e) pusiUinws (Chapuis) rettuipennis (School) shenefelti (Nllnberg) semisulmtus (Sehedl) suhahditus (Sch~dl) truncatkauda. (School) turgif1"ot18 (Sc:hedl) truncatigranuSflS (Selledl) umbrosus (Schl,,-dl) truncatipel1.niJI (Schedl) Myoplatypus DUlIlplatypus hipo"" (Blaodf(»-d) aetdi.dentalus (Muraymna) br"evicornis (Wood) lUhmcus (Chapuis) cormexus (Wood) agnutus (School) .flavicomis (FalJricius) alg"""" (SchedJ) prenexus (Wood) antiwC£p1wli (School) ,enerus (Wood) Iriu=as (Blandfo,d) calamus (Blandford) OropWtypus cavus (Strohmeyer) quad.ridenUUllS (Olivier) cAevrolati (Chapuis) cupula.d", (School) Platyphy= cupulatus (Chapuil:» cvnvexu.' (SchedL) decem (Sampson) laticoUis (Chapuis) fa1catus (Strohmeyer) oblWiWi (Chap\lis) fOlficul,(J (Chapuis) pouteri(l(; (Wood) hamatus (Blandford) oonfabri (Reichardt) lepidus (Chapuis) luniger (Motschulsky) MegaplatlJPUS malaisei (School) arlecarinablS (Scht',dl) m«ritimus (School) attenlus (School) nomUl{lanae (Browne) auricularis (Chapuis) omega (SchooL) auritus (Chapuis) pa/1idus (Chapuis) baksi (Chapuis) pmiperda (Schcdl) bicornis (Nunberl-:> pscudQCupulat", (School) bulens (Schcdl) ternlis (Murnyama) binocIul.uY (Chapuis) tentdssinms (School) brevicaudatw (Nunhery,) tetrl/.C{."rUS (Beeson) caraoonis (School) umhrou= (School) carinifer (SchedL) (Ulcinatm (Blandford) chiriquensi' (Wood) 280 GREAT BAS TN NATUHALIST [Volume 53

coru;iliatus (Schedl) raU£'Wi (Schedl) cun-sequens (Scheel!) reichei (Chapuis) contractus (Chapllis) robustus (Chapuis) costipennis (Scheel!) salvini (Blandford) c1wvidens (Scheel!) schmidti (Chapuis) darlit¥itoni (Reichardt) sexcost,atus (Chapuis) drmta.fus (Dalman) simplicifonnis (Wood) desultor (Schedl) sobrinus (Schedl) deymllei (Chapuis) sUaVifel" (SchcdJ) diductus (Chapuis) suboblitaratus (Schedl) discicoUis (ChaplIis) subsulcatus (Chapuis) dv,'coidalis (Schedl) tiriosensis (Reichardt) distinguendu,' (Scheell) tuben'1J.la,t11s (Chapuis) ddohratus (Blandf()rd) umbonatus (Blandford) durus (Schedl) w"sinus (Scheel!) egm{!,ui.s (Schedl) 1.1rS11S (Schcd!) elongatus (Chapllis) eq1uulorensis (Schedl) Euplatypu.~, exaratus (BlandfOrd) aequalicinctus (Scheel!) exitialis (Wood) alienus (Scheel!) exitioslis (Scheell) alt.ernans (Chapnis) flexiosus (Sched!) angustatulus (Wood) fossulatus (Chapuis) angustatu.~ (Chapuis) fragosus (Schcld) angustioris (Scheell) Juscus (Chapuis) araucariae (Scheel!) godmani (Blandford) areolatus (Sched!) granarius (Scheell) beUus (Schedl) gregalis (Scheel!) bilobatus (Strohmeyer) holdJw1lsi (ScheelJ) cornpositus (Say) ignotus (Sched!) contextus (Scheel!) imporcatus (Blandford) COroMtus (Schedl) insidiosus (Scheel!) costaricensiB (Schedl) insignatus (Scheel!) crihricollis (Blandford) irmiolat1/.s (Sdwdl) cuspidatus (Svhed!) irregtdaris (Scheell) deC01"WJ' (Schedl) ir'1"epel"tus (Sched!) dignat1.l.~ (Scheell) ifT1J{)tHs (Scl1edl) dimuliatus (Chapuis) jelskii (Nllnberg) dissimilis (Chapuis) konincki (Chapuis) dissipahi.lis (Scheell) lafertei (Chapllis) efferatus (Scheell) lal;reillei (ChaIHlis) haagi (Chapn!s) limhatus (Chapllis) hUJfI.,l' (Chapuis) limtl(,'US' (Wood) hintzi (Schaufuss) limtus (B!amH(wel) immunis (Sched!) luriclus (Chapllis) l,amiwltus (Schcdl) malignus (Scheell) longior (Wood) lnarginatlls (Chapuis) longius (Wood) Inlltatus (Chapllis) longulus (Chapnis) rUJ,vatTodeandmdei (MarcHi) mLulagascal"iensis (Chapuis) neglectus (Sehedl) minusculus (Schedl) nitidicoUis (Sched!) mulsanti (Chapnis) nudatus (Wood) otiosus (Sched!) ohliteratus (Hlanclf(}rd) parallelus (Fabricius) occipitalis (Chapnis) patmus (Chupnis) olivieri (Chapnis) pennirnicus (Schedl) perhinodulus (Schedl) pertusus (Chapuis) perrruwginatus (Scheell) pini (Hopkins) perm()d.{~stus (Scheel!) IJorosus (Hlandf()fd) pernudus(Sehedl) psctulolongulus (Sched!) peruanus (NunlJerg) pulicaris (Chapllis) p01Tectus (Chapllis) roberti (Chapilis) psefulodignatu',. (Sched!) rugosifi"ons (Schedl) pseudoplicntus (Schedl) sanf4cn.a:ensis (Mlltchlcr) quaesitu.R (Sched!) segnis (Chapuis) quinquecostatus (Chapnis) simplicijonnis 01'Iood) ram.ali (Scheell) sinuosus (Chapuis) 1993] REVISION OF PLATYPO()I\)A!i 281

mlutfls (Chapuis) com,:innulus (Blandford) slrintus (Chapuis) Cflixus (Schedl) tragus (School) excisus (Chapuis) tricu.,])idatus (Schedl) hwnilLs (Chu(>uic;) lrispinatulus (Schtxll) ;nocessus (Schedl) trispillatu.~ (Schedl) 111llrcidus (Blandford) truncalus (Chapuis) OnllJtus (School) MlIIts (Blandford) rmllidipetlnis (Blandford) percomis (Schelll) Baiocis perdiligell.' (Sdledl) See Wood & Bri~t cl992:1Z15-1217 ralzchurgi (Chupl.lis) slrialopenni,~ (Sdledl) Epiplatyptls suhilariu8 (Schedl) {/(lnexus (Schedl) ustulatus (Chapllis) mmexus 0""ood) (rpplarwlils (Wood) Cylin(lr01J(Jlpus bmslliensis (Nunberg) (See Wood & Bright d992:1217-12l9) cQmplanalus (Sdlt..'t.Il) deceplor (Wood) Tr;ozaslus deplanatus (Wood) Sec (Wood & Bright cl992:1219-1221) discolor (DIandfon:l) ettge,sttls (Wood) Me.